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BOSTON MEDICAL LIBRARY 

IN THE 

FRANCIS A. COUNTWAY 

LIBRARY OF MEDfCINE 




'jt"»»». ' 



THE 



JOURNAL 



or 



4NATOMY AND PHYSIOLOGY 



I 



CONDUOTKD BY 



G. M. HUMPHKY, ILD. F.R.S. 



or AVATOMT Of THE UXXTXBftITT OV CAMBUIMI. 
BOSOEABT FELLOW OF DOimKO CX>LLBOI: 

AVD 



WM. TURNER, M.R 

PmOPBBOB OP ASATOMT US TKB WXTTWUm OF IDIVBVBOH. 



VOLUME VL 

(SECOND SERIES, VOL. V.) 



MACMILLAN AND CO. 

eambt(fei8^ and loiOion. 
1872. 




PBINTBD BT a J. OLA.T, ILA. 
▲T THK UBIYBBSITT PBB88. 



f 



f 



Journal of ^natom^ m\f ^fi^siologp. 



^ THE MUSCLES AND NERVES OF THE CRYPTO- 
BRANCHUS JAPONlCtJS. By Pkofessor Humphry. 
(PL I. to IV.) 

Being so fortunate as to obtain the fine specimen of the great 
I Japanese Salamander which died recently from a gash across 
I its throat in the Zoological Gardens, in Regent's Park, and 
having the advantage of my friend and assistant Mr Anningson's 
k skill for its dissection, I felt that the opportunity of making a 
^ more careful examination of the muscles of the animal with 
their nerves than had hitherto been done was not to be lost. 
.Hjrrtl, in his Crypiobranchus Japonicua, 1865, does not describe 
either the muscles or the nerves, and Drs F. J. J. Schmidt, 
Q. J. Goddard, and van der Hoeven (Aanteekeningen ever de 
anatomie van den Cryptobranchus Japonicua, Haarlefa, 1862) do 
not give any account of the spinal nerves. 

The chief interest in the muscular system of this animal 

(and the other Urodelans) consists in the massing together of 

the several muscles, or rather the imperfect segmentation and 

invidualization of them ; so that it affords us, to some extent, a 

natural means of ascertaining the relationships of the muscles 

of the higher animals to one another, and so of grouping them. 

In instituting the comparisons requisite for this, it is difficult 

to exercise sufficient caution, and to avoid the errors which the 

necessities of nomenclature and the straining for over-exactness 

of homological interpretation are apt to induce. Especially is 

this the case if our views are warped by any theory we are dis- 

ym&A to &vour. Such influences work upon us imperceptibly ; 

and to be altogether free from them is more than is to be ex- 

/lected, even if it is to be desired. Facts will gravitate from 

tbeir chtum, and they do so in some minds more quickly, in some 

in better order than in others; and while we gain the good, we 

VOL. VI. 1 



2 i^ROFESSOR HUMPHRY. 

mtifjjtabicte by and beware of the evils of the tendency. I ex- 
press these thoughts becauae they have been present with me 
throughout the investigation of the Cryptobranch* 

FASCIAL INVESTMENT. 

The muscles of the trunk, tail, and limbs, are covered by an 
investment of fascia or tough areolar tissue, which is connected 
with the transverse intermuscular septa of the trunk and tail 
more closely than with the muscles, and from the deeper strata 
of which processes pass between the muscles, and are attached 
to the bones. One of these processes (Fig. 1) forming a sheet 
or lamina on either side, near the mesial line above, dips down- 
wards, covering the edge of the dorsal muscle, and meets its 
fellow of the opposite side along the line of the spinous processes 
of the vertebrae to which the two are attached. They thus en- 
close a triangular interval between them and the superficial 
stratum, which is continued across the middle line; and this 
interval contains fat, and affords a convenient passage for blood- 
vessels. A similar disposition exists along the ventral mesial 
line and along the lateral lines in the tail, as we shall find 
presently. Superficially it is connected with the skin and with 
the muscular fibres passing from point to point of the skin ; and^ 
which is very important to remark, it is closely connected with 
the trapezius, latimmua dorsi and the fore part of the pectorcUis, 
as well as with the suhcutaneus coUi, the depressor mandibtdcB, 
And the constrictor faticium, I say closely connected, because 
the fibres of these muscles are interwoven with and so take 
origin firom it, and in this respect differ from most or all of the 
other muscles of the trunk and limbs. In front the fascial in- 
vestment is continued dorsally, over the head to the margins of 
the upper jaw; and, ventrally, it is attached to the margin of 
the lower jaw. 

The interior of the trunk is lined by a similar investment 
(fascia transversaHs), somewhat more compact, though thinner 
than that on the exterior, which is connected with the trans- 
verse septa extending through the inner layer of the abdominal 
muscles. It is continued forwards to and along the base of 
the skull above, and, beneath, over (deeper than and attached 



THE CRYPTOBRANCHUS JAPONICUS. 8 

to) the branchial cartilages and the hyoid, to the inferior max-* 
ilia. Behind it forms a thick fascial stratum inside the pelvis, 
with which it is connected ; and it is continued, between the 
caudal ventral muscles on the two sides, to the haemal spines 
of the caudal vertebrsB, where it bleuds with the mesial ventral 
process of the external fascial investment\ 

Though thus described as an external and internal invest- 
ment, these fascial strata are parts of one fibrous system con- 
tinuous at the oral and anal ends of the animal, and connected 
with each other along the length of the animal by intervening 
septa — ^fibrous, cartilaginous, and osseous — ^which, though histo- 
logically varying, are members of the same system, and in the 
interspaces between which lie the muscles with their blood- 
vessels and nerves. 

In Fishes and in the Lepidosiren the external fascial invest- 
ment is, especially at the sides of the body, scarcely developed 
as a separate stratum, and the transverse intermuscular septa 
pass through it into the skin and bind the skin with its ap- 
pendages closely to the lateral muscle. Near the upper and 
lower edges and immediately over the lateral septum it is more 
free and more easily detached ; and there is frequently subcu- 
taneous muscle present at these parts. 

FASCIA AND MUSCLES OF TAIL. 

The tail is the simplest part of a vertebrate animal. It con- 
sifitB merely of axially disposed bony pieces or vertebrsB whioh 
are moveable upon one another, of the muscles which move 
them, of the nerves which excite and regulate the movements, 
and of the vessels which carry the nutritive materials to and 
fro. In Urodelans, as in Fishes, the movements of the bony 
pieces are restricted, or nearly so, to one plane —a vertical move- 
ment in a horizontal plane — and the disposition of the muscles 
is very simple. The muscles, indeed, composed of longitudinal 
parallel fibres, are arranged as in the Fish in two flat masses, 
one on either side of the tail, which, may be named after their 
homologues in the Fish, the * great lateral muscles.* They are 

1 These fascial inveBtmento, external and internal, are deecribed by Schmidt, 
Oeddard, and ran der Hoeyen, though not quite in same manner as above. 

1—2 



4 PROFESSOR HUMPHRY. 

separated from one another, that is, the lateral muscle of the 
one side is separated from that on the other by membranous 
septa passing vertically in the mesial line above and below 
from the vertebrsB ; and ossifications extending into these septa 
above and below constitute the ' neural' and 'haemal* spines. 

Each lateral muscle is, as in the Fish, divided into an upper 
or 'dorsal* and a lower or 'ventral' muscle by a horizontal 
membranous septum — ^which may be called the ' lateral septum' 
— stretched from the sides of the vertebrae nearly midway be- 
tween the upper and lower edges of the animal, and ossifica- 
tions extending into these septa on the two sides would con- 
stitute 'transverse' or 'lateral' processes. Such ossification 
however has not taken place except in the fore part of the tail. 

Each of the septa (Fig. 1) as it approacees the surface splits 
into two laminae which extend in opposite directions beneath 
the skin and meet and unite with the adjacent laminae of other 
septa Thus the laminae resulting from the splitting of the 
neural septum are spread out, right and left, upon the dorsal 
muscles, and meeting the ascending laminae from the lateral 
septa, form sheaths enclosing the dorsal muscles. The same 
is the case with the laminae of the ventral septum ; and the 
four muscles, the two ' dorsal' and the two ' ventral' of the 
two sides, are enclosed in sheaths, more or less distinct, formed 
by the laminae of the neural, ventral and lateral septa. 

Finally (Fig. 2), the four longitudinal caudal muscles — the 
two dorsal and the two ventral muscles — are divided trans- 
versely by septa^ extending with more or less obliquity back- 
wards or forwards from the intervals between the several 
vertebrae to the enclosing sheaths just described and the skin. 
So that each of the longitudinal muscles is transversely seg- 
mented into pieces corresponding with the several vertebrae. 
These transverse septa are tough and form as it were part of the 
muscle itself, being connecting media between the ends of the 
muscular fibres of the segments. They are not disposed in true 
vertical planes, but form slight curves with the convexity back- 
wards above and below the lateral septum ^ 

* Schmidt, Qoddard, and van der Hoeyen oonnt 46 of these tendinous in- 
■eriptions, 14 or 15 in the trunk, and the remainder in the tail, where the 
mnBcnlar intervals between them are narrower. 



THE CRYPTOBRANCHUS JAPONICUS. 5 

It is worth while to remark that the intervals or channels 
between the diverging laminse of the longitudinal septa are oc- 
cupied by fat, and serve as recesses for the passage of blood- 
vessels, which are thus arranged in four longitudinal trunks or 
systems — a ' dorsal/ a * ventral' and two * lateral/ The interval 
in the ventral septum is thicker than in the other parts, and 
the fat-mass occupying it is larger and extends in upon the 
stunted or bifurcated haemal spines. A transverse system of 
blood-vessels lies upon the transverse septa; and the several 
transverse vessels being connected with the four longitudinal 
trunks form communications between them, and discharge their 
blood into them. 

In addition to the four fat-masses disposed with the longitu- 
dinal series of vessels are four deep fat-masses lying beneath 
the four lateral muscles, and lying therefore above and beneath 
the transverse processes and lateral septa. The nerves, as they 
pass from between the vertebrae transversely to the muscles, 
cross superficially with regard to these deep fat-masses. 

The several fat-masses serve the purpose of giving size and 
lightness to the tail, making it a large and, by reason of the 
muscles, powerful propelling organ. Each attains its maximum 
thickness near the middle of the tail and diminishes towards 
the tip. 

The caudal muscles therefore consist of a ' dorsal' and ' ven- 
tral' muscle on each side, passing from behind forwards, and 
transversely segmented by the membranous septa and the verte- 
bral processes running into it. 

The several parts, muscular and other, of the animal above 
the level of the lateral septa present but little variety in differ- 
ent regions of the tail, or indeed in the different regions of the 
animal. Accordingly the dorsal muscle extends in the simple 
form I have described along the whole length of the tail and 
indeed onwards through the abdominal and thoracic regions to 
the head. 

The parts of the animal beneath the level of the lateral septa, 
on the contrary, undergo various and extensive modifications in 
consequence of the presence of the limbs and the abdominal 
thoracic and cervical viscera, which are all placed in the ventral 
division, that is in the division of the animal beneath the plane 



6 PROFESSOR HUMPHRY. 

of the transverse vertebral processes and the transverse or late- 
ral septa connected with those processes. The effect of these 
modifications is seen in the fore part of the ventral muscle of 
the tail. Behind and in the greater part of its extent it, 
like the dorsal muscle, is simple. Anteriorly, however, it is 
more or less interrupted and broken up by the proximity of 
the pelvis and hind limb. 

The upper portion of it encounters the ilium, which is deve- 
loped in the middle of the thickness of one of the transverse 
septa; and the middle stratum of this portion of the muscle 
is interrupted by or inserted into it and constitutes the iUo- 
cavdal muscle. The superficial stratum is continued forwards 
over the ilium, and being expanded, together with the rest of 
this part of the animal for the accommodation of the viscera, 
contributes to the formation of a layer of the abdominal muscles. 
The deepest stratum is continued forwards beneath the ilium, 
is similarly expanded, and joins the deepest stratum of the abdo- 
minal muscles. 

The undermost part of the ventral muscle, the part lying 
next to the ventral fat-mass, becomes separated from the re- 
mainder as it approaches the pelvis and hind limb, loses the 
transverse inscriptions, and divides into three. Of these one — 

* ischio-cavdalis* — situated nearest the middle line and the 
most superficial, arises from the extremities of the haemal spines 
and is inserted into the hindermost margin of the ischium^; 
a second division — ' caudo-cruralW — arises a little more late- 
rally from the haemal spines and arches and loses itself in 
the superficial stratum of the hinder part of a broad muscle 
(Figs. 8 and 9), which ia passing from the pelvic shield to the 
hind limb. It cuts this muscle nearly at right angles, and being 
inserted tendinous into it produces a tendinous inscription in it. 
A portion, however, of the caudo-crural does not lose itself in 
the broad muscle just mentioned, but is continued on along the 
hinder aspect of the thigh and leg into the musculo-tendinous 

1 It arises rather from the side of the ends of the haemal processes, being 
separated from the muscle of the opposite side by the fat-mass which lies upon 
(beneath) the middle parts of the extremities of the hsmal processes. 

The intervals between the hiemal spines are occupied by short antero-pbsterior 

* interspinons * muBcles, which form part of the ventral muscle, and the mar- 
ginal fibres of which are continued into the ischio-caudaUs, 



THE CBTFTOBBANCHUS JAPONICUS. 7 

mass of the sole, and so on to the extremities of the digits. It 
may be called ' caucUhpedcUts' and it constitutes a continuous 
muscle reachiug from the middle of the tail to the ends of the 
toes. Its fibres are partially interrupted by a tendinous in- 
scription, and it is joined beyond that point by a portion of the 
muscle arising from the ischium in which there is no tendinous 
inscription. (Fig. 10.) 

The third division — * caudO'/emorcUis' — arising somewhsut 
more laterally and deeply from the haemal arches, is inserted by 
a strong tendon into the middle of the under (plantar) surface 
of the femur. 

This disposition of the caudal muscles corresponds with that in 
the other Urodelans* (Axolotl, Menobranch and Newt) which I baye 
examined; the only difference being, that in these the upper and 
larger — the iliac — ^portion of the ventral muscle does not afford so thick 
a covering stratum to the ilium, and consequently that bone, occupy- 
ing as it does in Cryptobranch the plane of one of the intermuscular 
septa, approaches nearer to the suriaoa Internally the ilium is, as 
in Cryptobranch, lined by a deep stratum of the caudal muscle which 
is continued into the abdomen. 

The T&ANSV£RS6 INSCRIPTIONS or 'sclerotomes' which, in these 
animals, as in the fish, divide the lateral muscles into so many 
'myotomes,' serve the purpose of binding the muscles throughout 
their whole length and depth to the vertebral column, preventing 
their starting from the column towards the ai*c of the curve, or their 
having a tendency to do so, when the tail is bent to one side or the 
other under their contraction ; also by preventing the continuity of 
the muscular fibres, and by difiusing among many the force of the 
pull consequent on the contraction of any one, they add greatly to the 
strength of the whole. It is obvious that if each muscular fibre had 
been continued from end to end of the animal, and, further, had been 
required, as must be the case in these animals, and in fishes especially, 
to contract in its whole length simultaneously for the purpose of 
effecting those enei'getic violent flexures which produce the dart- 
ings and leapings of these animals, it would have been liable to 
rupture under its own force. The interruption however of the fibre 
by an inscription common to it with others diffuses the pull of the 
several parts of each fibre among many, and enables them all better 
to combine in . a simultaneous effort. The arrangement does not 
interfere with the nerve-supply because each myotome receives its 
own nerves from its own division of the spinal cord through its 
vertebral foramen, in addition to the filaments from the lateral 
nerve which travels along the lateral septum to the tail and then 

^ It oonesponds also generaUy with the aoooont giren by Mr Mivart of the 
^enopoma and Menobranob, Proc. Zool, Soe, April 22 and June 24, 1S69. 



8 PROFESSOR HUMPHRY. 

breaks up iuto branches (p. 47). When, however, as in the case 
of the caodo-femoral, caudo-crural, <fec., a part is segmented from 
the rest for the purpose of independent action requiring inde|)en- 
dent nerve-supply, then, partly for the better distribution of the 
nerves through its substance, the inscriptions disap{>ear, the con- 
tinuity of the muscular fibres being established through them so as 
to cause their obliteration. It is probable also, taking into account 
the peculiar vibratory or successional manner of action of the several 
pirts of a fibre by which sustained contraction is effected, that con- 
tinued action may have relation to continuity of independent fibres ; 
whereas violent sudden efforts are associated with interruption of 
fibres, or, as in the case of the heart, with interlacing and intercom- 
munication of fibres, which would have much the same effect. Prac- 
tically, at any rate, we find where sustained action is required the 
muscular fibres are parallel and uninterrupted, but where sudden 
violent efforts are needed the fibres are interlaced and communicating, 
or are segmented by transverse tendinous inscriptions. 

It is obvious that just as the extension of the muscular fibres 
through or over the tendinous inscriptionB would cause fusion or 
ankylosis of the myotomes, and the occurrence of this partially and 
in vaned ways will lead to varied dispositions and divisions and 
complications of the muscular structure: so the extension of the 
tendinous inscriptions through or over the myotomes would cause 
fusion or ankylosis of the sclerotomes; and tlie ossification of the 
thus extended sclerotomes may lead to varied prolongations and com- 
plications of the bony skeleton. 

I have mentioned iucidently that the limb-girdles and the limbs 
are placed beneath the lateral septa in connection with the ventral 
parte of the lateral muscles and have little or no relation to the 
dorsal parts, that the same is the case with the ribs, and that the iliac 
bones are, like the ribs (pp. 6 and 10), the result of ossifications in the 
ventral tranh verse intermuscular septa. It does not hence follow that 
the iliac bones are precisely the serial homologues of the vertebral 
ribs or ^pleurapophyses.' An objection to that view is presented by 
the fact that there intervenes in the Cryptobranch and other Urode- 
lans, between the iliac bones and the vertebral transverse processes on 
either eide, a distinct bone which must correspond with a rib, and 
which in Menobranch has the elongated characters of a rib. This, 
together with the ventro-mesial position and relations of the pelvic 
bones and their freedom from the vertebral column in some animals 
(Whales, Snakes and Lepidosiren), indicate a serial correspondence 
with the skeletal formations in the sternal rather than with those 
in the vertebral ('pleurapophysial') region of the visceral wall. I 
avoid applying the term 'hsemapophysiar to the former because I 
think it by no means clear that the visceral cavity and its wall are to 
be regarded as identical with the hiemal canal and its wall. Indeed, as 
stated above, the lining of the visceral cavity (fascia tramversalia) in 
the Cryptobranch is continued from the interior of the pelvis beneath 
the haemal arches of the tail where it blends with the subhsdnud 
septum ; and the position which a backward or caudal prolongation of 



THE CBYFTOBRANCHTTS JAPONICUS. 9 

the Tiacend cavity would occupj appears to be that oocnpied bj 
the ventral or sabhjemal CEit-masa. If this view is correct the visceral 
wall represents a stratum superficial to that of the hiemal wall; and 
the hemapophyses, instead of being in the same plane and correspond- 
ing with the lower parts of the visceral wall, or, instead of being 
expanded to the level of those parts — the pelvis, linea alba, sternum 
and sternal ribs, shoulder-girdle, and hyoid appai*atus — are rudimentaiy 
or abortive in the visceral region. The representatives of them are 
to be sought in the fibrous structure covering the aorta and its 
branches, and lining the crura of the diaphragm, not in the subvis- 
ceral part of the wall of the viceral cavity. 



THE MUSCLES OF THE TRUNEL 

The muacles of the trunk may clearly be regarded as, in 
part at least, formed by an extension of the caudal muscles 
forwards. Like these they are divided by a 'lateral septum' 
into a 'dorsal' and a 'ventral' series; and the tenner present 
very little difference from those in the tail. The septum is 
continued from the extremity of the tail forwards, beneath the 
latissimtu dorsi and the muscles passing from the dorsum of the 
head and spine to the scapula and over the neck, as far as the 
head. The fibrous sheet which forms it slants from the ex- 
terior somewhat upwards to the upper surface of the ribs and 
transverse processes, so that the dorsal series of muscles overlaps 
the ventral to a slight extent: and owing to the greater ex- 
pansion of the inferior or ventral part of the trunk, in com- 
parison with that of the upper or dorsal part, the lateral septum 
is here situated nearer to the dorsal than to the ventral edge 
of the animal. 

The TRANSVERSE SEPTA, a Serial continuation of those in the 
tail, are directed from the median line above, at first very ob- 
liquely outwards and backwards through the inner part of the 
dorsal muscles ; gradually, as they pass over the ribs they be- 
come less oblique, describing a curve with the convexity back- 
wards, and near the lateral septum they are almost transverse. 
Beneath the lateral septum they are continued with slight ob- 
liquity backwards, and then again become transverse, thus 
forming in the ventral region a gentle curve with the con- 
vexity backwards, like that which is seen in the dorsal and in 
the ventral muscle of the tail. The direction therefore is not 



10 PROFESSOR UUICPHRY. 

80 zigzag as in the fi^, or quite so much so as represented by 
van der Hoeven. Viewed from the interior of the trunk they 
are seen to be directed from the mesial line, on either side, 
over the several intervertebral substances, obliquely backwards 
and outwards, upon the bodies of the vertebrae and over the 
ribs, in a direction corresponding with that of the transverse 
septa on the corresponding part of the exterior. Indeed they 
are a continuation of them through the thickness of the animaL 
They are easily traced down to the ribs, and the ribs appear to 
be the result of ossification occurring in them, beneath the plane 
of the lateral septum, that is, in the ventral parts of them. Be» 
yond the extremities of the ribs the direction is rather suddenly 
changed, becoming more transverse; and towards the middle 
line they are, as on the exterior, quite transverse. (Fig. 4.) 

Direction of the Muscular Fibres. 

In the DORSAL series (above the lateral septum) the direc- 
tion of the muscular fibres is parallel with the axis of the ani- 
mal. As in the tail, they simply pass between the transverse 
intermuscular septa and between the vertebral processes, and 
^how no distinct indications of grouping into distinct bundles or 
muscles. They are moreover not interrupted by the presence 
of the limb-girdles. They require therefore no special de- 
scription. 

In the VENTRAL series (beneath the lateral septum) however 
the case is different. The intermuscular septa preserve their 
serial order, that is, follow one another in regular succession, as 
in the dorsal aeries, and are continued through the entire thick- 
ness of the muscular wall of the abdomen, and the muscular 
fibres pass from septum to septum, being limited to the inter- 
vals between the septa; but the muscular wall is expanded to 
cover the abdoi)ainal cavity and accommodate the viscera, and it 
is interrupted, or partially interrupted, by the limb-girdles; 
moreover, the direction which the muscular fibres take between 
the septa varies in different jxurts of the abdominal wall, and at 
different d&pths of the same part The difference in direction 
9X different depths causes a more or less distinct division into 
planer; and the difference in direction at different parts of the 



THE CBYPTOBBANCHrS JAP0NICU8. 11 

abdonuBal wall causes a more or less distinct division into 
wctora. 

Near the exterior the muscular fibres are directed, between 
the transverse intermuscular septa, from before backwards, and 
ventrally towards the linea alba. This disposition distinguishes 
them from those of the subjacent layer which take a different 
direction, and gives rise to a stratum constituting an * Miquua 
ejrtemus' muscle. The obliquity is most marked near the lateral 
septum. At a distance from that it gradually diminishes, and 
near the mesial line the fibres acquire an antero-posterior or 
straight direction, and constitute a superficial layer of the 
* rectus abdominiaJ 

In the stratum next subjacent the fibres are directed frx)m 
before backwards, and dorsally towards the lateral septum, and 
thus is constituted an 'obliquus intemua' The obliquity is here 
also more marked near the lateral septum diminishing towards 
the mesial line; and the fibres here acquiring an antero-pos- 
terior or straight direction constitute a deeper layer of the 
' rectus' 

The 'rectus abdominis' is therefore simply the resultant of 
the altered direction of the fibres of the internal and external 
oblique, and consists of two layers which may, to some extent, 
be separated from each other, and which are continuous respec- 
tively with the external and internal oblique muscles\ (Fig. 7.) 

In the deepest stratum of all the muscular fibres with their 
intermuscular septa extend, under the surface of the bodies of 
the vertebrae^ as far as the middle line. In this situation they 
are directed antero-posteriorly or straight, and constitute what 
may be designated a * subvertebral rectusV More laterally, 
where they lie beneath the ribs and are connected, by means 

^ It would be more striotlj correct to say that the obliqui are the resultant 
of AD altered direction of the fibres of the recttu^ and the two layers which they 
form are consequent on the difference in the direction which their fibres re- 
speotively take in their variation from the antero-posterior course of those of the 
rectuB. 

This fusion of the external and internal oblique muscles with the rectus, 
which I have indicated diagrammatioally in Fig. 7, was pointed out by Mr Mivart 
in his description of the Menopom% Proc. ZooL Soc., April 22, lS6tf, p. 268, i^ 
well as in that of the Iguana, Proc. ZooL Soc., 1867, p. 770.- 

s This muscle is desorilrad as rectus trunci internus by Schmidt, Goddard, 
and van der Hoeven ; but they do not mention its connection with the traiuver^ 
talis. By Biivart it is regarded, in Menopoma (Proc. ZooU Soc, April 22, 1862, 
pi 260), as part of the retrahens eottarum* 



12 PROFESSOR HUMPHRY. 

of the intermuscular septa, with the ribs, they acquire a slant 
from behind outwards as well as forwards, and constitute a series 
of ^depressores costarum! More laterally still the obliquity 
increases to an almost transverse direction. This part of the 
stratum is the ' transversalis abdominis.* Towards the linea 
alba it degenerates into a fascial layer, which is continued 
above (on the deeper a£pect of) the rectus abdominis to join 
its fellow in the middle line. 

The nerve-trunks lie in the chief part of their course behind 
the intermuscular septa, between the internal and the middle 
muscular planes, t.e. between the subvertebral rectus, the depres- 
sores costarum and the transversalis, on the one side, and the 
internal oblique on the other. (Fig. 4.) 

It will thus be perceived that the muscular fibres lying 
nearest to the mesial plane of the animal, ventral as well as 
dorsal, are directed ant^ro-posteriorly, corresponding with the 
axis of the trunk, and constitute so many pairs o{* recti,* 'supra- 
vertebral,' ' sub- vertebral,' and 'sub-abdominal;' whereas the 
fibres more laterally situated are oblique, and constitute obliqui 
and transversi. This accords with the directions in which the 
muscular forces are required to operate in the mesial and lateral 
planes. 

It will be perceived further that the several abdominal mus- 
cles are essentially one, transversely segmented, muscular mass, 
that the stratification or segmentation into planes is simply the 
result of the variation in the direction of the fibres at different 
depths, and that the recti differ from the obliqui and transversi 
only in the circumstance that this variation in direction at dif- 
ferent depths has not occurred in and near the mesial plane of 
the animal where they are situated. In the Fish where the 
variation in direction does not commonly take place at any part, 
not being required by the form and movements of the animal^ 
the ' rectus* character pervades the whole of the ventral muscular 
system of the trunk, and there are no obliqui and transversi. 

Lastly, the stratification, even in the lateral parts of the 
Cryptobranch and other Urodelans is incomplete, intermediate 
between the condition of the Fish and that of the higher verte- 
brates^ fon^much as it affects only the muscular fibres, and does 
not extend to the intermuscular septa. In Birds and Mammals, 



THE CRYPTOBRANCHUS JAP0NICU8. 13 

tvhere more independent action, not only of the several strata^ 
but of the several parts of each stratum, may be presumed to 
take place, the stratification is more complete, and the septa 
have in great measure disappeared Still even in them the sub- 
abdominal rectus retains commonly more or less of the original 
septal and unstratified type. The persistence of these characters 
in it contrasting so markedly with the alterations which have 
been wrought in the more lateral muscles, and the consequent 
complete separation of the one from the other, mask the natural 
relations of the several abdominal muscles, and make us over- 
look the fact that they are all derivatives from one simple 
antero-posteriorly disposed muscular mass. 

Descbiftion of the Abdominal Muscles. 

The obliquus extemus (Figs. 2, 3, 5, 7) may be described 
as arising from the extremities and outer surfaces of the ribs 
from the second to that just in front of the pelvis, inclusive, by 
digitations, of which the foremost are very distinct. Further 
backwards the digitations are less distinct, and the muscular 
fibres arise pai-tly from the lateral septum as well as appa- 
rently, to some extent, from the fascial tissue covering the dorsal 
muscles ; this however is not quite clear. Joined by a more 
superficial portion from the ventral surface of the thorax \ it 
extends over the side of the abdomen to the middle line and 
the edge of the prepubic shield and cornu. The fibres, for the 
most part oblique, are near the middle line antero-posteriorly 
disposed, blended with, or constituting the superficial layer of 
the rectus. The hinder part of the muscle, dorsally, is not 
very defined, being partly attached to the ilium and partly 
lost in &scial tissue : ventrally it is attached to the edge of 
the prepubic shield and cornu superficial to the pyramtdo' 
lis: and it is strongly attached to the spinous or marsupial 
tubercle which projects forwards from the hinder part of the 
edge of the prepubic shield. It is also continued into the 
gracilis, which arises firom the under surface of the prepubic 
shield and pubes; and it is attached to the ala of the ilium, 
while some of the fibres passing over it are lost in fascial 
tissue ; and between the ilium and the gracilis or pubes it is 

^ This is the pectoral xnaMle, Me p. 83. 



14 PROFESSOB HUMPHRY. 

continued into the fascial tissue, which spreads over the dorsal 
aspect of the thigh, knee and leg. A deeper portion extends 
from between the anterior edge of the ilium and the pubic 
spine as a fibrous process, which acquires close connection with 
the hip-joint. (Fig. 2. e, o\) 

If it is traced in the opposite direction, it may be described 
as radiating from the sides of the prepubic cartilage and the 
pubes, anteriorly and dorsally. The fibres nearest the middle 
line run forwards as rectus, having a slight connection with the 
sternum, and are continued beneath, i.e. superficial to the coracoid 
of the same side, forming the superficial thoracic portion men- 
tioned above. Some of these are lost upon the fascial tissue of 
the coracoid ; but the greater number diverge laterally and are 
attached to the humerus, contributing largely to the formation 
of the pectoralis major. The fibres of the oblique situated more 
laterally than these take a deeper course, above the coracoid, 
and are attached to the extremities of the ribs as far forward 
as the second. Thus the fore or under part of the scapular arch 
i(the hinder portion of the coracoid) projects back between the 
' recto-pectorcUis* and the 'obliquus' divisions of the oblique 
muscle. The fibres of the oblique arising still more laterally, 
are directed with increasing obliquity upwards to the ribs, the 
lateral septum, and the dorsal fascia, and backwards to the ala 
of the ilium and the fascial tissue of the thigh. 

It will be understood that this, which we call a single 
muscle, is composed of successional series of fibres or myotomes 
passing from rib to rib and from transverse line to transverse 
line, the transverse lines being continuous with the ribs. 

Obliquus internus. (Figs. 2, 4, 5, 7.) If the fibres of the 
external oblique are cot through, the subjacent sti*atum of 
fibres taking a different direction, viz. from behind forwards 
and {fiwards and constituting the obUquiLs irUemus, is disclosed 
without difficulty, forasmuch as the change in direction takes 
place abruptly. 

The internal oblique is more massive than the external 
oblique, and more obviously a continuation of the iliac part of 
the caudal ventral muscle. Indeed that muscle, or great part 
of it, is continued forwards into the internal oblique, the ala of 
the ilium, which constitutes the notable line of demarcation 



THE CBTPTOBRAirCHTTS JAPOKICUS. 15 

between the two, not being seen on the exterior or the interior, 
and being, as before said, apparently an ossification in the thick- 
ness of the intermascolar septum. 

The internal oblique may be described therefore as arising 
from the upper part of the ala of the ilium, from the iliac part 
of the caudal ventral muscle passing above and beneath the 
ala of the ilium, and from the several ribs^ as &r forwards 
as the fourth, in a deeper plane than the external oblique. 
Its origin does not however extend so far forwaids by two 
ribs as that of the external oblique The fibres are, in the 
lateral psirt of the animal, directed obliquely from behind for* 
wards and inwards, or mesially between the intermuscular 
septa ; but near the ventral mesial line they take a more 
antero-posterior direction, constituting the deep stratum of 
the rectus. The hindmost fibres in this manner join or con- 
tribute to the rectus, in front of the prepubic cartilage and 
comu, and are therefore not attached directly to the pubes. 
Anteriorly it is continued beneath the abdomen and thorax 
into the neck passing above, that is deeper than, the coracoid 
to the branchial cartilages and the hyoid. Its mesial partr-* 
the part forming the deeper stratum of the rectus — when 
passing above the coracoid comes into contact with and 
is attached to the deeper plate of the sternum', the edge of 
the plate being received among its fibres; and between thife 
point and the hyoid apparatus, it forms the sterno-hyoid. As it 
travels forward iu the neck it is joined, near the hyoid, by a 
narrow muscle from the anterior edge of the scapula, which is 
the cmo-hycid. (Fig. 5, o. h,) Or, the relations of the omo-hyoid 
to the internal oblique may be more clearly expressed by stating 
that as the latter muscle proceeds backwards from the hyoid 
a portion is detached to, or acquires an attachment to, the fore 
part of the scapula and forms the omo-hyoid. 

The hinder and inferior margin of the internal oblique pass- 

^ The ftenram (Fig. 5) is a yexy rndimentaiy ti tru ct ure , tonsisting of a 
broad heaii-shaped plate of cartilage, with the rounded apex directed forwaids ; 
and it is deeply notched or bleft on either tide anteriorly for the reception of 
the edgee of the large ooracoids. So de^ly ia it cleft that it might be regarded 
aa oomriating of two plates, a superficial and a deeper, blended in the middle 
fine and behind. It is the deeper plate which is involyed at its margin in the 
jntemal obliqne. The clefts on the two sides are not quite in the same plane, 
that of the left side extending a liUle beneath that on the right side. This 
permits the edge of the toft ooraooid to pass superficially to or beneath the ri|^. 



16 PROFESSOR HUMPHRY. 

ing from the ilium to form the deeper stratum of the rectus, 
is tolerably defined ; and there is an interval between it and 
the retiring angle of the pubes and prepubic shield, which is 
occupied by the Pj/ramidaUs. (Kg. 2. Py.) The fibres of this 
muscle, arising from the anterior edge of the pubic shield and 
the tubercle or spine ^ which bounds that edge laterally, pass 
forwards and inwards to the edge of the prepubic cartilage and 
oomu, and some of them are continued further forwards, blend- 
ing with the edge of the rectus. It lies between the external 
obUque and the transversaUs, its outer margin being parallel 
with, though at a short distance from, the inner margin of the 
internal oblique. Its fibres take the same direction as those 
of the internal oblique and are in the same plane with them 
and appear to belong to the same series. In the Axolotl this 
is proved to be the case by the continuity between the two, 
that is, by the absence of the interval which separates them in 
the Cryptobranch. 

The Transversalis (Fig. 4) is connected with the ribs and 
the transverse intermuscular septa in a still deeper plane. Near 
the vertebral column the direction of its fibres is nearly antero- 
posterior, and they are with difiiculty distinguished from those 
of the internal oblique ; still the division between the two is 
marked by the passage of the nerves. More externally their slant 
first resembles then exceeds that of the fibres of the internal 
oblique (from behind forwards and inwards), and finally they 
become almost transverse. At about an inch and half from the 
mesial line, at a part that is con*esponding with the formation 
of the rectus in the manner I have described, the muscular 

1 The relation of the mosoles shows this lateral prepnbie tubercle to be the 
homologae of the spine of the pabes in man, and of the marsupial bone in mar- 
supials. I have dissected ^e muscles of this part carefully in the Wombat, Par- 
aeyon, and Echidna, and find that the external obUque fibres forming the outer 
oolumns of the external ring are inserted into the outer edge of the marsupial 
bone, whereas the fibres forming the internal column pass across to the base of 
the marsupial bone of the opposite side and interlace with the corresponding 
fibres of the opposite side. The pyramidaHt muscle, which is large in most of 
these animals, arises from the inner edge of the marsupial bone, and extends 
forwards and inwards in front of the rectus and behind the external oblique to 
the middle line, becoming blended with the rectus posteriorly so as not to be 
easily distinguished from it. In these animals, with the exception of the Wom- 
bat, the internal oblique has no connection with the marsupial bone and the 
pyramidalis, but passes deeper than they, and deeper than the rectus. In the 
Wombat a few of tiie hindmost fibres of the internal oblique are connected with 
tlie marsupial bone. 



THE CBTPTOB&ANGHUS JAFONICUa I7 

fibres of ihe ircmsversalis end in a tendinous sheet which is con- 
tinued above (deeper than) the rectus, to join its fellow in the 
middle line. Anteriorly the tranwersaUs takes origin from the 
ribs as far forwards as the internal oblique ; and it forms a thin 
sheet passing above the coraeoids and the sternum. It is sepa- 
rated from the coraeoids by the internal oblique ; but it comes 
into contact with the deeper surface of the sternum near the 
mesial line. A space is here left uncovered in consequence of 
the fibres of the internal oblique being limited to the margin of 
the deeper plate of the sternum, and the transver$aii$ accord- 
ingly is attached here. In front of this it blends with the 
internal oblique or is lost in areolar tissue. Towards the 
hinder part of the abdomen it is inserted into the upper surface 
of the prepubic cartilage and comu in a manner corresponding 
with its attachment to the upper surface of the sternum ; and 
it is traceable within the pelvis in company with the deepest 
stratum of the internal oblique into connection with the caudal 
muscle. 

Where the fibres of this deepest or transveraalis stratum of 
the abdominal muscles lie beneath the ribs passing from rib 
to rib, directly or through the medium of the intermuscular 
septa in which the ribs lie, and passing from the sides of the 
vertebral bodies to the ribs, they constitute a series of depres* 
sores costarwm (Fig. 4, i?, C)\ and where they lie beneath the 
bodies of the vertebrae they constitute what I have called a 
sUbvertAral rectus. In the former situation (where they form 
the depressores costamm) they slant from behind forwards and 
outwards like the fibres of the internal oblique; but in the 
latter situation (where they form the subvertebrcU rectus) they 
take a more antero-posterior direction. The subvertebral rectus 
is continued forwards beneath the bodies of the cervical ver- 
tebrae and is attached to the skull, so constituting a longus coUi 
and rectus capitis. It will be understood that there is no line 
of demarcation between these several muscles, the whole — 
trcuisversaliSy d^ressores eostarum, subvertebral rectus, longus 
colli, rectus capitis — ^being one continuous sheet of muscular 
fibres, with transverse septa or inscriptions, extending beneath 
the vertebral column, forwards, as far as the head, backwards, 
into the caudal muscles, and continued laterally beneath the 

VOL, VI. 2 



18 PBOFESSOB HUHPHBT, 

internal oblique to the ribs, the ventral mesial line, the pre- 
pubic cartilage, and the sternum. 

Mesially, beneath the vertebral bodies, the muscle is sepa- 
rated from that of the opposite side by an interval in T?hich 
lie longitudinal subvertebral vessels. Branches from these ves- 
sels pass transversely between the several vertebral bodies and 
the subvertebral rectus supplying both, and are continued as 
'intercostal' and 'lumbar' vessels in company with the nerves, 
behind the intermuscular septa, between the transversalis and 
internal oblique. 

There can, I think, be little doubt that the cnira of the dtaphrctgm 
in Mammals are formed by the lumbar parts of the subvertebral 
reetue bending downwards on the sides of the aorta and encircling 
it, and that the btteral parts of the diaphragm are in like manner 
formed by the iDflaction of the lateral parts — the depressores costartuah 
fCnd ira/neverealis parts — of the same sheet. 

A series of muscular fibres extending forwards to the head 
in the plane of, and taking the same antero-posterior direction 
as, those of the external and internal oblique near the vertebral 
column and forming a continuation of them, constitute the 
acaleni and the recti laterales. They pass between the trans- 
verse processes, between the ribs and between the intermuscular 
septa, and are separated from the rectus capitis (the anterior 
part of the subvei*tebral rectus) by the emerging nerves. 



MUSCLES OF THE HIND LIMB. 

The muscles of the hind limb are, in accordance with the 
chief movements of the several parts of the limb, arranged in 
two series, one upon the plantar and the other upon the dorsal 
aspect of the limb; and in the furrows between the two, along 
the anterior or tibial and the posterior or fibular edges, are 
contained the chief blood-vessels and nerves of the limb. This 
at least is the case in the thigh; in the leg and foot they lie 
more near the middle of the dorsal and plantar aspects of the 
limb between the superficial and the deep strata of the 
muscles. 



THE GBTPTOBRAHCHUS JAFONICUS. 19 

PiAKTAB Muscles or Thigh. 

In the thigh the plantar mass arises from both surfaces — 
the npper or abdominal and the inferior surfaces— of the under 
or sub-costal parts of the broad pelvic shield, that is, from the 
region of the pelvis situated on the ventral or plantar side of 
the hip-joint, and consisting of the ischium and the pubes. The 
mass is continued anteriorly into the Miqtio^ecttis muscle of 
the abdomen, and posteriorly is connected with the caudo- 
crural muscle. It thus intervenes between and connects (or 
may be regarded as an extension of) the mesial portions of the 
ventral muscles of the abdomen and the tail 

It is divided into two STRATA. Of these the SUPEBFICIAL 
is a large, broad, unsegmented, and thick mass. It arises from 
near the mesial line of the large pelvic shield in nearly its 
whole length, from the insertion of the ischto-caudal, behind, to 
the external oblique, with which it is blended, in front. The 
grreater part of it (all of it except the fibres presently to be 
mentioned as joining the catulo-pedat) is inserted into the 
upper two-thirds of the anterior {tibial) edge of the tibia in- 
clining to the plantar aspect It is the great flexor and ad- 
ductor muscle of the leg, and corresponds with the gracilis^ 
semitendirumts and semimernbranostis. About one-third from 
its origin it is joined nearly at right angles by the fibres of the 
caud(h€rural (p. 6, Fig. 8), which, or the greater number of 
them, terminate, tendinous, in its hinder and superficial part, 
causing a tendinous inscription in it. 

The hinder part in which the caudo-crural thus terminates 
appears to correspond with the iemilendinanu, the anterior superfi- 
cial part being referable to the gracUi$ and the deeper part to the 
temimembranotua ; and I cannot but sospect that the inscription thus 
formed may afibrd a more probable explanation than has yet been 
given of the remarkable inscription in the gemUendinogus of man. If 
it be so it is a curious illustration of how &r back in the animal 
series we may search for the explanation of peculiarities in the human 
fiame, and, further, of the fact that minor features may appear or 
reappear and be the only trace of those conditions in the general plan 
of development which seem to have been their cause ; for I need 
scarcely say that of the ecmdo<rural muscle itself there is no trace in 



A bundle of the fibres of the hinder part of the muscular 
mass we are considering, unmarked by an inscriptioti, joins a 

2—2 



20 PROFESSOR HUMPHRT. 

similar bundle. ftom the ca/udo-crural^ which is to some extent 
marked by an inscription, and they together form the cmudo^ 
pedal (Figs. 3 and 10). This runs down the back of the leg 
and expands into the fascia of the leg and sole, reaching to the 
extremity of the digita There is thus one continuous muscle 
extending from the middle of the tail to the ends of the digits, 
deriving occasional fibres in its course from the pelvis, and, as 
ire shall find, fi*om the femur, the leg-bones, and the tarsus. 

The DEEPER STRATUM of the plantar mass is in three parts. 
A middle part arises beneath (deeper than) the preceding from 
the under surface of the pelvic shield. It is large, covers the 
obturator bole, and receives the nerve em^ging from that 
hole. It is inserted into the tibial side and middle line (Jinea 
aspera) of the plantsM^ surface of the femur and into a line 
passing from the Imea aspera to the tibial condyle. It repre- 
sents the addti^tors and the extemai obturator. A hinder — 
ischio-femoral — ^p^ arising from the hinder edge of the ischium 
is inserted rather above and behind the preceding into the pro- 
jection on the fibular side of the upper end of the femur. It 
represents the quadratvs femaris, the ffemeUi, and the internal 
obturator. It is separated from the contiguous edge of the 
dorsal muscular mass, or the ilio-femoral portion of it, by the 
sciatic nerve, which lies between them, as it passes from the in- 
terior of the pelvis. An anterior — supra-pubic — ^portion repre- 
senting the pectineus (Fig. 2, Feet-) arises from the whole of the 
upper or abdominal surface of the pubes, behind the pubic 
spine and the pyramidalis, not therefore from the pr^ubic car- 
tilage. It here extends as far as the middle line, meeting its 
fellow of the opposite side. It is a large muscle, passes over 
the anterior edge of the pubes and the inner part of the hip- 
joint, and is inserted into the linea aspera and tibial side c^ 
the plantar surface of the femur in conjunction with the mid- 
dle portion. Moreover, it expands over the dorsaX surface of 
the lower third of the femur, immediately above the knee-joint. 
It here extends round the fibular side of the femur, as far as 
the plantar or popliteal surface, and therefore yearly embraoes 
the lower part of the shaft of the femur. Where it passes over 
the fore part of the pubes it is separated from the middle, or 
adductor, portion of its stratum by a prolongation of the tendon 



THE CETrrOBKAirCHUB JMFQBICUS. 21 

of ike eKternal oblique which extends from the spiae of the 
pabes, aloag the Anterior maigin of the pubes, to the hip-joint ^ 

DoBSAL Muscles of Thioh. 

The dorsal muscular mass is, aboYe, connected with the 
supra-cozal part of the pelvis, i.e. with the ilium. Below^ it is 
partly connected with the femur and partly extends upon the 
leg. This causes a division, as in the plantar mass, into a 

ST7PERFICIAL or ILIO-CBURAL and a DEEP or ILIO-FEMORAL 
STRATUlL 

The SUPERFICIAL STRATUM is divided into three long bands 
or sectors. The inner (Fig. 2, i2, t), arising by a flat tendon 
from the pelvis over the hip-joint, external to the pectineuSy 
descends over the anterior or tibial part of the knee close to 
the joint, and passing beneath the origin of the tibialis ariHcus 
from the femur is inserted into the dorsal surface of the head of 
the tibia^ dose to the knee. I name this recttu femaris irUer- 
ntis, and judge that it corresponds with the so-named muscle in 
Birds. It has no distinct representative in mammals, though 
its insertion nearly resembles that of the aartorius. It acts as ^ 
an extensor of the leg. 

* ThU k not the Ugamentum leret. Indeed, I do aot find lo dietiBet a re* 
preeentailTe of that ligament either in the hip or the shoulder as figured and 
deseribed by Hyrtl (Tab. y. fig. 8, and Tab. tx. fig. 1). In the femur and the 
humerus the cartilage forms a broad horse-shoe belt over the somewhat flattened 
upper end of each ; and into the ooneavity of the oresoent on either side, which 
ia thua left uneorex^d bgr cartilage, the capeole of the joint ascends hinder than 
at other parts of the circumference of the bone. The fibrous tissue of the cap- 
aide 18 aUo rather thick at these parts and so forms ridges, as it were, projecting 
into the joint, and passing from the head of the bone upwards and downwards, 
k> the ilium and to the ischium in the case of the hip, and to the seapula and to 
the oonuxnd in the ease of the shoulder. Soppose the ends of the cartilage 
horse-shoe to be prolonged in either instanoe and to meet forming a circle^ 
endowing a central dimple, and cutting off from the rest of the capsule the pro- 
jecting portion of it inserted into t^t dimple, a veritable ligamantum tere$ 
would be formed. This might take place either above or below. It does so 
below, towards the isdiium, oommonly in the hip ; and in the shoulder, a pro- 
jection of the capsule forming the gleno-hmnexal ligament towards the upper or 
scapular side presents an approach to the same thing, and is a reminder of the 
pacts in the (hTptofaraneh. In the Jouim. of AntU, it. 28, 1 have called atten- 
tion to a recess and dimple in the astragalus of Unau and of AL They are 
samilar to these in the humerus and femur of Gryptobranch ; and with these are 
aasoffisted ia eaeh animal a similar appipaoh to the formation of a ligamentum 
teres from a part of the posterior peroneo-tarsal ligament which ascends into the 
jMMs sad dimple. 

If this is, as I belieTe, the true explanation of the Ugamentum teres^ we must 
regard that ligament as merely a detached portion of the capsule of the joint, 
<iid not, as has been soggmted, the representative of a tendon or other stmstuie* 



22 PROFESSOB' HUMPHBT. 

The middle sector (Fig. 2, GL 2) arises from the outer sor* 
face of the ilium, half an inch from the hip-joint, and descends 
as a broad strap-like muscle over the thigh and over the middle 
of the knee, superficial to the inner sector and the origin of 
the dorsal leg-muscles from the femur. It does not acquire 
any attachment to the upper end of the tibia, being separated 
from it and from the knee by the dorsal muscles of the leg, 
extending up to the femur ; but it expands over those muscles 
and sends a process deeper, between the tibiaiia anticus and 
the extensor digit&rum, which is attached to the lower end of 
the tibia and expands upon the ankle. I name this part of 
the ilio-crural stratum, 'gliUeo rectus^ believing it to represent 
the gluteus maximus (if there is any representative of that 
muscle in Urodelans) and the rectus femoris of mammals. 

The outer sector (Fig. 2, B) arises from the ilium just 
beneath the preceding, and so close to it that it may almost be 
said to arise by a common tendon vrith it ; it descends along 
the fibular side of the thigh and terminates in a broad tendon 
which, insinuating itself between the peroneus muscle and the 
fibula, is inserted into the upper third of the shaft of that 
bone. The peroneal nerve (Fig. 10, P) crossing beneath it, runs 
along its upper edge to the dorsum of the leg. The relation of 
the nerve, as well as the insertion of the muscle, indicate this 
outer or ilio-fibular sector to be the biceps jlexor cruris; and it 
afibrds an example, not uncommon, of a muscle which belongs to 
the dorsal or extensor series becoming, by virtue of its position 
upon the side of a joint, an adjunct in its action to the plantar 
or flexor group. It is accompanied, in the distal part of its 
course, by a distinct spindle-shaped muscle (Fig. 10, F^f), which 
arises, by a tendon, from the plantar aspect of the femur, just 
beneath the insertion of the tendon of the caudo-femond, and 
is inserted tendinous into the middle third of the shaft of the 
fibula beneath the tendon of the biceps. This femoro-fibular 
muscle appears to represent the short or femoral origin of the 
biceps. 

The DEEPER or ilio-femoral stratum of the dorsal mass 
covers the iliac part of the hip-joint. It arises from the outer and 
anterior surface of the ilium near the joint, between the pectineuB 
anteriorly and internally and the ischio-femoral posteriorly and 



THE CRTPTOBEANCHU8 JAPONICUS. 23 

externally, being separated irom the former by the origin of 
the iniemal redus and from the i9Ghuhfemoral by the sciatic 
nerve. It moreover extends a little upon the internal surface 
of the ilium, and still further upon the adjacent internal surface 
of the ischium, where it reaches behind the pectineua as far as 
ihe middle line. It is inserted into the fibular side of the middle 
of the Unea asperOy close to the ccMdo-femaral and the femoro- 
fibular and close to the adductors, which are inserted into the 
tibial side of that line. Its attachment extends also from the 
linea aspera upon the fibular side of the plantar surface of the 
femur. Its origin from the ilium extends on either side of, or 
rather, in front and behind the origin of the gluteo-rectus and 
biceps: and the part in front of those muscles, lying upon the 
front of the hip and the ilium, appears to correspond with the 
iUacus intemua, while the part lying behind those muscles and 
lying upon the back of the hip and the ilium appears to corre- 
i3>ond with the lesser glutei and the pyri/ormis. 

It must be remarked that the ilium is chiefly occupied by the 
aitachmentB of the caudal and abdominal musoles — the internal 
oblique more particularly — and little space is left for the glutei. 
They are accordingly almost abortive. The gluteus maaamus is 
absent^ unless it is, as I have sttppoeed probable, blended in the part 
of the ilio-crural section which I call gUiieo-rectas ; and the other 
glutei are very small ; and in order to gain sufficient space for attach- 
ment they extend upon the inner surface of the ilium, and still 
more upon that of the ischium, thus spreading beyond their proper 
area and encroaching upon the territory appertaining to the plantar 
mupcles, more particularly occupying the ground from which the 
internal obturator usually arises. This, at least, I suppose to be the 
case. It must however be stated that the muscles immediately 
surrounding a ball-and-socket joint in such an animal as the Crypto- 
branch somewhat resemble the capsule of the joint itself in the 
mode in which they invest it^ and in the imperfection of their seg* 
mentation ; so that the division into separate muscles is rather ar- 
bitrary, and the nomenclature proportionately uncertain. 

It will have been remarked that the deeper parts of the 
extensor cruris, viz. the vasti and cmreuSf which are such con* 
stant elements in higher animals, do not exist in the Crypto- 
branch, and the lower space of the femur usually devoted to 
their origin is occupied by the insertion of the pectineits, while 
the upper part does not give attachment to any muscle. 

The dorsal aspect of the thigh is covered by areolar or 



24 PROFESSOR HUHPHBT. 

fascial tissue in addition to the musdes above enumerated;* 
and this is an extension of or is continuous with the hinder 
portion of the external oblique between the ala of the ilium 
and the spine of the pubes* 

. Plaktab Muscles of Leq and Foot. 

Below the knee the plantar aspect of the limb is occupied by 
a broad thick pronato-flexor mass, extending from the femur 
to the ends of the toes, and with some difficulty distinguishable 
into parts — strata and sectors. It arises from the plantar sur- 
face of the fibular side of the limb, namely, from the fibular 
condyle of the femur, from the fibula in its whole length and 
the fibular side of the tarsus, being continuous with the ahduc- 
tor minini digiU. Superficially, it is blended with the cauio^ 
pedal, extends over the sole and divides into five tendons, 
which pass to the terminal phalanges of the five digits. De- 
tachments from the deeper surfaces of these tendons pass to 
the first and second phalanges; and still deeper parts of the 
muscle are inserted into, as well as arise from, the proximal 
and distal row of tarsal bones and the metatarsals. The chief 
direction of the fibres is from the fibular towards the tibial 
side of the limb as well as downwards. Near the surface the 
fibres are nearly vertical and have, in the main, a flexor action. 
Traced more deeply they are, for the most part, more oblique ; 
and the deepest of all are transverse and act simply as pronators. 

The mass is partially divided into a SUPERFICIAL and a 
DEEP STRATUM, with the chief nerves of the back of the leg 
and sole running between them. The two strata are however 
UendiBd together above and still more below. 

The SUPERFICIAL STRATUM (Figs. 10 and 11), which is the 
part blended with the caudo-pedal, extends from the fibular 
condyle and the fibula to the ends of the digits. Its fibres 
have in the main a vertical direction, and exercise in the main 
a flexor function. It corresponds apparently with the gastro^ 
cnemius, solevs and plantar fascia^ and with the plantaris and 
yfaror breffia digitorum (the last two should be regarded as one, 
forming a flexor sublimia digitorum^). I will recur to the dis- 

^ The terms *#uMtmu* and 'profundus* are greatly to be preferred to 
* brevU* and * lon(fua*in destgnating theM flexor w^flf M? teft of the digUs ; beoaniHi^ 



THB CRTF10B&ANCHU8 JAPOVICUS. 25 

poation of its diyisioiis in the digits after descrildng tbe fi m or 

profwdiis. 

The BEEP STRATUM is disposed ia an interesting manner. Its 
fibres are more oblique and have more of a pronator function 
than those of the superficial stratum. It is composed, first and 
chiefly, 61 a mass of fibres (Fig. II, Pr. p) derived, slightly, from 
the fibular condyle of the femur in conjunction with those of 
the superficial stratum, but, chiefly, from the fibula and the 
fibular part of the tarsus — the two fibular tarsal bones — and 
slightly from the adjacent tarsal bones. They are directed 
downwards and inwards to the distal row of tarsal bones, to the 
metatarsal of digit I. and slightly to the metatarsals of li. and 
IIL, and also join the deeper surface of those divisions of the 
superficial stratum which pass to the flexor tendons of digits 
I. II. and in. This mass may be called * pronator pedis* ^ : it 
appears to combine the representatives of the tibialis posHcus 
and those portions of the flexor digitarum profundus which we 
call the flexor longus poWids and the acoessorius. 

A second part of the deep STRATtnc (Fig. II, Fl, pr.) still 
lying beneath the nerves is a long, thin muscle, quite separate 
in its upper part^ which arises from the upper end of the fibula^ 
descends upon the main n^ass of the deep stratum (the pro- 
nator pedis), the fibres of which cross obliquely from the fibida 
beneath it. Having reached the tarsus it expauds, trumpet* 
like, beneath it and divides into five muscular bundles to the 
five digits. I have said that it lies, in the leg and in the 
proximal part of the tarsus, superficial to the promUor pedis; 
but its insertion into the digits is deeper ; and in order to reach 
this deeper plane it, or rather the part of it destined to digits 
I. n. and m., curls round the fibular edge of the part of the 
pronator pedis passing to the flswor subUmis, so as to come into 

in oonjnnetion with the pUmtarU, whioh is reaUy the upper or omral part of it, 
the JUxor iublimU {brevU usually termed in man) is quite as long or longer 
than ihejlexar^fundut; and the relation oi the flexor subHmU to the olonkrrtf 
-^« cardinal pomt in the anatomy of the muscles of the leg illustrated in most 
mammalH — is ignored or rather concealed by the term *br€vu* of human 
anatomists. 

^ I call the muscle * pronator pedis ^ to facilitate subsequent descriptions, and 
beeaose the name indicates an important part of its function ; but even in this 
animal it has a flexor action, especially the fibres of it joining the flexor sub' 
RmU, and its representatives in the higher animals, in which pronation and 
anpiftatfoa ol Ihe foot do not take plaao, haTe a flexor action only. 



26 PBOFESSOR HUUPHBT. 

close proximity with the tarsus. As it exiMmds beneath' the 
distal row of tarsal cartilages to reach the digits on the tibial 
side, it crosses the insertion, of the 'proftialUyr pedis into those 
cartilages, and itself acquires a connection with the cartilages, 
sending some fibres to them and deriving some fibres from 
them, and some of its fibres are here blended with those of the 
pronator pedis. 

In Menopoma (Fig. 13) the interraption by the projecting tarsal 
cartilage is more complete than in Cryptobranch ; so that the long 
band descending the leg is inserted there, or nearly so, and short 
muscles to the digits ta^e their origin there. It seems not impro- 
bable that the flexor profundus is compounded of detachments from 
short flexor muscles arising from the tarsus with a detachment 
from the proncUor pedis inserted into the tarsus ; and may it not 
be that the fibrous mass, or sesamoid ossicle so commonly found in 
the tendon of this flexor and in its homologue of the fpre limb, is a 
representative of the skeletal structure which in these aniTnals in- 
tervenes between the upper and the lower parts of the muscle, and 
which becomes detached with the muscular fibres when the continuity 
of the two parts of the muscle is established f 

Further, are not sesamoid ossicles in other parts similar detach- 
ments finom the bones near which they lie? 

Each of its five digital bundles subdivides into three, a 
middle and somewhat superficial part which is inserted into 
the base of the proximal phalanx, and two lateral parts which 
are attached to the sides of the metacarpal. The subdivision, 
which passes to the fibular side of met. v. is thin; and the 
division which passes to the poUex subdivides only into two, 
one to the middle and the other to the fibular side of the proxi- 
mal phalanx. This muscle represents in the main that portion 
of iheflsxor dtgitorum which we usually call the flexor longus 
digitorum. I say in the main, because the representatives of 
the two parts of tlus stratum, which I have designated pronator 
pedis and flexor profimdus digitonmi, are, in other animala, 
variously blended to make up the flexor haUuds, fleosor lonffus 
digitcrum, and the a4Xsessor%us. 

The arrangement of each of the divisions of the flexor subL 
digitorvm resembles generally that of the divisions of the^/Iftror 
profimdus digitorum just described. Each (Fig. 11, a) sub* 
divides into three, of which the two lateral pass to the sides of 
the approximated ends of the metacarpal bone and the prozi* 



THS CBTFT0BRAKCHU8 JAPONICUB. 27 

mal phalanx, while the middle portion passes on to the second 
or t^minal phalaooLx. That at least is the case in digits I. il« 
and v., which have only two phalanges. In digits in. and iy.^ 
which have three phalanges, the middle portion again sub- 
divides into three, of which the lateral parts are attached to 
the sides of the penultimate phalanges, while the middle parts 
go onwards to be inserted into the terminal phalanges. So 
that digits I. IL and T. have the same complement of muscles 
as ni. and iv., but the second tripartite division does not take 
place in them. 

There are in the individual digits some exceptions to this 
arrangement which should be mentioned. Thus, in digit i. the 
tendon sends onlj one offset to the middle of the proximal 
phalanx, and then passes on to the terminal phalanx. In IIL 
the penultimate phalanx is treated like the proximal in I. re- 
ceiving only one ofiset to its middle from the tendon on its 
way to the terminal phalanx. In v. the tendon detaches the 
nsaal lateral ofibets to the approximated ends of the metacarpal 
and the proximal phalanx, and subsequently detaches an ofiset 
to the middle of the same phalanx before its insertion into the 
base of the terminal phalanx. 

These points are deserving of special notice, because the lateral 
offiets are probably the representatives of the lumbricales^ and the 
median offsets are probably the representatives of the retinactUa not 
unfrequently found in Mammals, and usaally observable in oonneo* 
tion with flexor tendons of the toes in Birds. 

The third part of the deep stratum (Fig. 14, Pr. t) lies be- 
neath, deeper than, the others, and is separate from them and 
composed of fibres still more obliquely, indeed almost trans- 
versely, directed. They form a square muscle, passing from the 
shaft of the fibula across the interosseous space to the shaft of 
the tibia. It may be called 'pronator tibice.^ The upper fibres^ 
have a slight inclination from the fibula downwards, while the 

^ Some ol the uppermost fibres are attached quite to the top of the fibula. 
In Bdxie the upper piurt, slightly separate from the rest, arises by a tendon from 
a aeeamoid behind the knee common to it with the descending tendon of the 
eando-femoral and the two strata of the fiexor mass; and through the medium of 
this sesamoid and the fibrous bands that unite it with the condyle, the upper 
part of the pronator Hbia deriTes its origin from the fibular condyle, and corre- 
aponds with the popliteu$ of mammals. 



26 PBOFESSOB HUMPHEY. 

lower fibres have a slight inclmation iipwahis, crossing b^ihd 
the upper. 

Tarso-meiatarsdlei (Fig. 12) are a aeries of short thick mus- 
cles passing from the under surface of the distal row of tarsal 
bones to the under surface of the metatarsals near the base. 
Each, like the flexors of the digits, divides into three, but the 
lateral portions arte rather more prolonged upon the metatarsals 
than are the middle portions. In digit I. the middle portion is 
wanting, and the laterals are very delicate In digit V. the di- 
vision into three parts is scarcely to be made out. 

MetacarpO'phaiangei (Fig. 12) arise from the under surface 
of the nietacarpals near the insertion of the middle portions of 
the tarso-metatarsals, and in some instances are continuous with 
them. They are present in all the digits^ and are ins^ted into 
the bases of the proximal phalanges, dose to, and blended with, 
the tendons <^ theflsxar digitorum longua inserted at the same 
parts. 

Phodangei (Fig. 12) are jn^esent only in digits lu. and lY. 
(the digits with three phalanges). They arise firom the under 
aurface of the proximal phaknges, and are inserted into the 
bases of the second phalanges blending with the psffts of the 
tendons of the flexor digitorum suhlimia which are attached 
here. There is a slight thickening in the tendons as they pass 
imder the joints suggestive of a sesamoid body. 

In higher animals the phalcmgei are absent, and the irUerosaei 
(p. 30), the tarso-mekUarsaUs, and the metatarso-phalangei are blended 
in the short flexors and interosaei. 

Addvctor minimi digiti is a piece of the flexor mass extend- 
ing from the lower end of the fibula to the fibular side of the 
tarsus and the base of met. v. 

DoBBAL Muscles of Leg and Foot. 

The muscles on the dorsal aspect of the leg and foot form a 
supinato-extensor mass corresponding antagonistically with the 
pronato*flexor mass on the plantar aspect, and, like it, consisi- 
ing of a superficial and a deep stratum. There is however st 
less amount of muscular substance, and less complexity in its 
disposition and subdivision. 



THE GBTPTOBBANCHUS JAPONICUS. SS 

The siTPBBnciAL STfiATam aiises by one broad tendon from 
the dorsal soi&ce of the fibular condyle of the femur. It soon 
diyides into three parts or Bectors — a 'tibial,' a ' fibular/ and an 
'intermediate/ 

The tibial sector — Hb4aU$ antioua (Fig. 15, T. cl) — is in- 
■erted along nearly ihe whole of the {ore part of the tibia. Its 
more superficial fibres extend over the ankle, and are inserted 
into the dorsal surfaoe of the proximal tarsal bone on tiiie tibial 

The deeper fibres of the intermediate sector — exi^ruor Umgnig 
digttorum (Fig. 15, K d) — are united to ana inserted with those 
of the preceding into the tibia^ The more superficial portion 
divides over the ankle into five broad tendons which reach the 
terminal phalanges of the five digits, and send down detach- 
ments from their deeper surface to the proximal phalanges. 
Moreover at each notch between the divisions into the five ten- 
dons a process passes into the interval between the subjacent 
XDetacarpals, and bifurcating extends a short distance along 
their contiguous sides and is inserted into them, so that traction 
of the muscle has the effect of approximating the digits. 

A delicate detachment (Fig. 15, E. d'.) from the fibular side 
of the muscle in close apposition to a similar detachment 
from the fibular sector, forming a peroneua tertius, is inserted 
into the proximal tarsal bone on the fibular side. 

The outer or fibular sector — -peroneus — is inserted into the 
fibula in nearly its whole length, with the exception of the 
slip just mentioned, which contributes to form the peroneus ter- 
titu. This muscle is therefore, in the Cryptobranch, derived 
from both the peroneus and the exteviaor digitorum. Some few 
of the fibres of the peroneus, it should be added^ extend over 
the end of the fibula, and are continuous with the fibres of the 
abductor minimi digiti. 

The DEEP STBATUic exists as a separate layer only at the 
lower part of the leg and on the foot. It consists of two parts. 

(1) Supinator pedis (Fig. 15, S, p.), a narrow band which 
arises from the dorsal surface of the lower end of the fibula, and 
crosses the ankle and tarsus obliquely to the base of the tibial 
aide g[ the metatarsal bone of digit n. (Digit i. is absent in the 
hind limb,) 



30 PBOFESSbR HtTMPHRT. 

(2) Extensor brevis digkorum arises from the dorsal storfSace 
of the taiBUSy more particularly on the fibular side, and is in- 
serted into the under surface of the extensor tendons as they 
pass over the digits. A slip detaches itself from the tibial side 
of this muscle and joins the supinator pedis. 

Inierossei metacarpales occupy the proximal parts of the 
intervals between the metacarpals, passing nearly transversely 
between the adjacent sides of the several bone& Their free 
edges are defined and curved or semilunar, the fibres descending 
a little along the sides of the metacarpals, but they do not reach 
the phalanges. 



MUSCLES OF THE FORE LIMB. 

The fibro-cellular or fascial tissue which covers the dorsal 
muscles and extends upon the ventral muscles and upon the 
head is thicker over the dorsal muscles than elsewhere, and is 
especially thick near the head. It here furnishes attachment or 
origin to muscles passing upon the scapula and the throat 

There are four muscular sheets thus arising placed beneath 
one another and distinct from each other. The most superficial 
{subcutaneua coUi, Fig. 5, 8. c. C), the thinnest and least defined, 
is a cutaneous muscle the representative of the panniculus car» 
nasiM and the platyama myoides. Behind, it reaches as fax as 
the laiisaimua dorsi. Anteriorly and ventrally it meets itR 
fellow in the middle line, and is attached along the inner side 
of the edge of the lower jaw. Near the angle of the jaw its 
deeper surfieu^ is joined by the fibres of the myho-hyoid radiat- 
ing above it. Under the 8ubcutaneu8 colli is the depressor fnanr- 
dibulxB and then the consiricior fawsiv/m. 

The deepest is the trapezius (Fig. 6, 2V.), which arises 
from the fascia covering the dorsal muscle opposite the sca- 
pula and as far forward as the skull, also from the occipital bone 
above the insertion of the dorsal muscle, and round the side of 
that insertion from the lateral part of the occipital. This last 
part of its ori^ is very deep, reaching with the lateral septum 
to the base of the skull, and probably represents the cemco- 
humeral of certain mammals^ The fibres converge to be in* 



THE CEYFTOBRANGHU8 JAPONICUS. 31 

serted into the anterior edge of the scapula^ the upper edge 
of the preooracoid, and the retiring angle beween the scapula 
and the precoracoid. 

Levator gcapuUs (Fig. 6, L. sc.) is a long narrow muscle 
arising by a delicate tendon from the base of the skull close to 
the insertion of the subvertebral rectus, indeed looking like a 
derivation from it, and inserted into the middle of the outer 
surface of the suprascapula above the origin of the dorsalia 
scapula. 

It arises from the skull in this animal and the other TJrodelans 
probably, because of the proximity of the scapula to the head. It 
is a very definite muscle in them, and attached only to snd near 
the angle of Ihe scapula. In Saurians^ where the davide reaches 
this angle, the muscle creeps upon it forming a levaiar davicul4»; 
and rudiments of the same are sometimes met with in Man and 
other mammals. 

SerraJtus magnus (Fig. 4, 8.) arises by two bundles from the 
fore parts of the ends of the second and third ribs, the two fore- 
most origins of the external oblique being from the hinder parts 
of the ends of those ribe. It is inserted into the under surface 
of the upper and also of the anterior part of the supra-scapula. 

The amo-hyoid (Fig. 6, 0. A.) passes from the anterior eflge of 
the scapula beneath the trapezius to the side of the ventral 
muscle advancing forwards to the hyoid, with which it is 
blended*. 

These are the four muscles which pass from the trunk to the 
scapula. 

The muscles of the fore limb, like those of the hind limb, 
may be classed as flexor or 'palmar/ and extensor or 'dorsal;' 
and, after the manner of the hind limb, of those which pass 
fit>m the shoulder-girdle to the limb, the palmar muscles take 
their origin from the coracoids, or subglenoid part of the girdle, 
and the dorsal muscles arise from the scapular or supraglenoid 
part of the girdle. Here, however, as in the hind limb, the 
muscles are not bound within rigid barriers, frmctional or terri- 
torial, but may be found to stray or glide more less across their 

■ In Kenobranoh the long preooiaooid eartilage lies upon the omo-hjoid, and 
Ibia mnsele ao it passes back beneath it divides into two, one portion going over 
the anterior, and the other over the jeK)sterior, edgs of the base of the preooraeoid 
to its outer surface, where the^ are inserted. 



32 PROFBSSOR HUBCPHRY. 

frontier lines into other districts, and sometimes pass from an 
extensor to a flexor office. 

On the Palmar Aspect of the Girdle akd Ark. 

The pectoral (Fig. 5, P.), is derived almost entirely from 
the superficial stratum of the obliquo-rectus of the abdomen, 
and is continuous with it. Where the fibres diverge from 
the trunk-muscle and take their independent course over the 
coracoid and to the humerus, they lose the tendinous inscrip- 
tions, tie. in the last four inches of their course. A few fibres 
are derived from the sternum; and as the muscle passes over 
the hinder part of the coracoid it acquires some addition from 
muscular fibres which arise from the coracoid. It is inserted 
into the radial edge of the radial tubercle near the upper end of 
the humerus, none of the fibres extending beyond this tubercle. 
Its anterior edge is continuous with fascial tissue superficial to 
the precoracoid and extending over the n6ck\ 

The coracoid and |»recoracoid cartilages in this animal are 
very large and form a considerable part of the glenoid cupj and 
the muscles arising from them are numerous, almost surround 
the head of humerus, and are difficult of interpretation. They 
arise chiefly from the outer' surface or the edge, and are as 
follows. 

1. A broad thin muscle, arising from the outer surface of 
the sternal or epicoracoid edge of the coracoid superficial to the 
htceps. It crosses the muscular fibres of the biceps superficially 
and transversely and converges to be inserted into the summit 
of the upper part of the radial tubercle of the humerus, just 
above the pectoral. It may be called eptcoraco-humeral (Kg. 
17)*. Some of its superficial fibres are blended with those 
of the under surface of the pectoral, and it is not improbably 
the representative of the pectoraUs minor of mammals. 

^ ThiB is the lascial inTestment already spoken of ^p. 3) as oorering tke 
body. Sohmidt, Goddard, and van der Hoevaa, speak of uie oonneetion of the 
pectoral with the sternum and ooraeoid, bnt do not mention its relati<»i to the 
external obliqne, or to the fascial tissue. 

* It ooiresponds, I think, with that deeoribed nndev this name in flie 
Echidna, Vy Mivart. Trant. Linn. Soe, zxr. 888. 



THE CKYPTOBRANCHUS JAPONICUS. 83 

2. Precor<io<hbrachud (Fig. 17, P. c. 6.), arises from the whole 
of the outer surface of the precoracoid cartilage, vrith the excep- 
tion of the marginal part. It lies in the same plane with the 
epicoraco-humeral, indeed is almost continuous with it, and is 
inserted, in close connection with it and with the pectoral, into 
the radial side of the uppermost part of the radial tubercle of 
the humerus*. 

3. Coraco'hrachialis longus (Figs. 17 and 18, (7. 6. Z.) is the 
largest of the muscles arising from the coracoid. It arises from 
the hinder edge of the coracoid and divides into two portions. 
Of these, the larger and inner or lower division is inserted into 
the ulnar edge of the humerus for a quarter of an inch above 
the internal condyle : the other division, being nearly fus large, 
is partly inserted into the side of the long tendon of the biceps, 
while a bundle of its fibres is continued on over the elbow, 
and is inserted into the ulna near the joint. 

This last-described division must represent the short or coracoid 
origin of the biceps in Man. There is no trace of it in Menobranch 
Axolotl or Newt. The muscle in them though large is confined to 
the humeruB in its insertion. 

4. Coraco-hrachicdts brevis (Fig. 17, C. h, brj) arises from 
the coracoid close to the shoulder-joint, between the preceding 
muscle and the joint, and also from the external surface of the 
hinder part of the coracoid near the joint. It passes beneath 
the biceps to the ulnar side of the humerus near the shoulder- 
joint, and to the base of the ulnar side of the radial tubercle of 
the humerus beneath the coraco-brachialis superficialis. It cor- 
responds with the ordinary mammalian coraco-brachial. 

The median nerve passes between these last two ; while the ninar 
vessel, and in animals where it is [>reseDt the ulnar uerve also, con- 
tinues its course behind both. 

1 This muBele is called subclavitu by Miyart, in his desoription of Menopoma 
and Menobranch, Proc. Zool. Soc. 1869, pp. 265 and 460, thongh he regards it 
as tlie same as that named by him epicoraeo-humeral in ^e Echidna. The re- 
lations of the mnsde to the mammalian mbelavitu do not appear to be snffi- 
eientiy dear to induce me to follow in the application of that name to it ; uid it 
Bjisee from the precoracoid rather than from the epicoracoid part of the girdle. 
In Menobranch it is inserted into the summit of the radial tnberole, and its 
under snrfaee is blended with the snpra-soapolar which makes its appearance in 
that animal though quite absent in Oryptobranch« 

VOL. VI. 3 



9i PBOFESSOR HUMFHBT. 

6. GoriiCQ'brcuJiialis quartus (jntbsoapularis) aiiseB from the 
mar^n of the coracoid and also of the scapula forming ihe 
inner edge of the glenoid cup, extending from the origin of the 
coraco-brachiah's brevts to and over that of the triceps and also 
a very short distance upon the adjacent inner surface of the 
scapula. It separates the origin of the coraco-olecranalts from 
the capsule and from the origin of the triceps. The latter 
expands somewhat into the capsule, and so separates this muscle 
hehind from the capsule. With that exception it is in imme- 
diate contact with the capsule which is thin under it. It is 
inserted into the upper part of the humerus behind the pre- 
ceding, that is, just behind the ulnar edge between the corcuxh 
brachialis brevts and the inner or third origin of the triceps. 
Indeed some of its fibres are continous with the former of those 
muscles in front, and with the latter behind. 

This mascle spreading itself to a greater extent upon the internal 
surface of the scapula in some animals becomes the subscapular. It 
is called subscapular by Mivart in Menopoma and Menobran<^ In 
the latter animal its extension upon the inner surface of the scapula 
is rather more decided than in Gryptobranch. 

6. CarOrCo-olecranalia arises, by a tendinous band, fix)m the 
edge of the coracoid, near the joint, internal to the middle of 
the preceding muscle, which separates it from the joint. It 
soon becomes muscular, descends the ulnar side of the arm dor- 
sally, joins the triceps and is inserted with it into the inner side 
of the olecranon. 

This coracoid accession to the triceps is common in Reptiles as 
well as in Urodelans. It results from the large relative size of the cora- 
coid, aud reminds us of the accession which tlie qttadrie^ extensor 
cruris in these animals receives in the form of what I have called w- 
terTial rectus from the inner side of the front of the hip-joint. Only, 
in consequence of the difference in the rotation of the two limbs, the 
extensor of the forearm being directed backwards, gains its accession 
from the back of the shoulder-girdle, while the extensor of the leg being 
turned forwards gains its accession ft»om the front of the pelvic girdle. 

The coraco-olecranalis is also an example of the deviation of a 
member of the flexor peries to an extensor function : just as the 
biceps fleocor cruris is an instance of the deviation of a member of the 
extensor series to the flexor function, the office in each instance bein^ 
determined by the position and attachment. 

7. Coraco-radialis, or biceps (Fig. 17, -B.), arises from the 



THE CBYPTOBRANCHUS JAPONICUS. 85 

external surface of the coracoid, between the epicoraco-brachial 
(pect, minor) and the short coraco-brachial, as a fan-shaped 
muscle, the fibres of which pass across the short coraco>brachial 
and soon converge into a long tendon, which runs down beneath 
the pectoral Having passed the pectoral it receives the fibres 
of the long coraco-brachial, passes over the elbow-joint, and is 
inserted into the palmar surface of the upper end of the radius 
close to the joint. It is supplied by the nerve which perfo-. 
rates the scapula and which supplies also the superficial coraco^ 
brachial 

I find the biceps corresponding with the above and distinct from 
the hraciiicdia anticus in Menobrancb, Newt, and Axolotl. In the 
Newt the long thin tendon is more closely related to the hrachialis 
anlicus, but does not seem to derive muscular fibres from it. As 
above mentioned, it does not in any of these animals derive fibres 
from -the coraco-brctckioMa langus. 

The Brachialis anticus arises from the radial side of the 
radial tubercle of the humerus, close, but on the opposite side 
of the tubercle, to the insertion of the pectoral, and beneath the 
insertion of the dorsalis scajmlce^. It passes along the shaft of 
the humerus, deriving no fibres from it, passes over the elbow 
superficial to the hicepa tendon, and is inserted about equally 
into the radius and ulna. Its foremost fibres are inserted into 
the ulnar side of the radius at a short distance from the joint, 
nearly in the situation corresponding with the middle of the 
insertion of the supinator longus in ourselve& Its hinder half, 
passing between this radial insertion and the insertion of the 
bic^y is attached to the adjacent palmar surface of the ulna. 



On the Dorsal Aspect of the Girdle and Arm. 

The trapezius already described, (p. 30). 

Latissimus dorsi arises from the fascial tissue over the dor* 
sal muscle, extending, forwards, nearly to the head, overlying the 
hinder part of the trapezius and, backwards, half way to the 
pelvis. It is a thin muscle and its edge is not very defined in 

> I. may observe that neither in this nor in the hind limb is the surfaoe for 
attacfainent of nraseles increased by intermascolar septa extending from the 
flidfis ol the homenu and femu; 

3—2 



36 PROFESSOR HUMPHRY. 

front or behind. The fibres converge and are blended with tli^ 
upper part of the scapular origin of the triceps. Through the 
medium of it some are connect'ed with the hinder maigin of the 
scapula near the glenoid cavity; but it has no direct attach- 
ment to the scapula^ and we cannot trace any of its fibres to the 
humerus. 

In these animals, in Keptiles and Birds, the lot, d, overlaps the 
irapeziua; the reverse being the case in mammals. 

Dor salts scapulas (Fig. 17, D. s.) arises from the outer ex- 
panded surface of the supra-scapula. It is a long trumpet- 
shaped muscle, and is inserted into the radial side of the radial 
tubercle of the humerus beneath the insertion of the pre* 
corac(hhrachiaL 

This is in Oryptobranch the only muscle, with the exception of 
the few fibres of the svhscapuUvris above mentioned, passing from the 
scapula to the humerus. In Menobranch there is in addition a 
muscle, blended with the deeper part of the precoraoo-brachial, which 
represents the guprc^-seapuUir, The darsalis scapulcB must represent^ 
in the main, the infrck-spiriatiu. Its superficial part^ probably, be- 
comes in other animals developed so as to contribute to the scapular 
part of the ddtoidy and its hinder part is segmented as the teres 
fninor. The davietdar part of the deltoid corresponds probably, to some 
extent, with the superficial fibres of the preoaraeo-braehial. Thus ihe 
clavicular and scapular parts of the deltoid come into relation with 
the trapeziiLS, which is, in Urodelans, inserted into both preooracoid 
and scapula. They overlie the supra- and irhfirO'Spinatus and teres 
minor musdefl, and have insertion corresponding with that of the pre- 
coraco-brachial and dorsaUs scapuke. The two parts are not uncom- 
monly separate. 

The few fibres of the svhsoapidaris constitute the only appearanoe 
of muscle upon the whole of the large extent of the concave under 
surface of the coraooids and scapula, so that the surface is free to play 
upon the convex outer surface of the ventral muscle on which it lies^ 

Triceps arises by a tendon from the posterior edge of the 
scapula, immediately behind the joint and in close connection 
with the capsule, which indeed it strengthens by expanding 
upon it. It soon becomes muscular and is joined by the 
latissimus dorsi, A second head arises from the radial side of 
the humerus at the base of the tubercle, and a third from the 
hinder and ulnar aspect of the humerus. These all unite; and 
the -muscle is inserted into the olecranon, being joined by the 
coraco'olecrnnah's. 



THE CBTFTOBRANCHTJS JAPOKICUS. S7 

» 

Palmab Muscles of Fokearm and Hand*. 

On the palmar aspect of the forearm the muscles form, as on 
the plantar aspect of the leg, a pronato-flexor mass, which is 
partially divided into two strata. 

The SUPERFICIAL STRATUM (Fig. 18) is in three sectors. A 
radial sector representing the Pronator teres and the Flexor 
carpi radialisy large and thick, passes from the anterior or pal- 
mar part of the ulnar condyle, commencing close to the joint 
and extending a little way up the humerus above the condyle, 
crosses the forearm obliquely, and is inserted into an oblique 
line on the radius, which passes across its palmar surface from 
the ulnar to the radial edge, and into the radial edge as far as 
the wrist, also into the radial side of the two radial cartilages 
of the wrist. The upper edge of its insertion is immediately 
below that of the hrachialie arvticua. The deepest portion of 
the muscle, arising from the condyle close to the joint and pass- 
ing to the ulnar edge of the radius, that is, to the upper part 
of the oblique line just mentioned, is in close contact with the 
pronator quadratue and at its insertion is blended with it, but 
is separated from it nearer the origin by the nerve (ulnar nerve) 
passing from the median nerve to the ulnar side of the forearm. 
This deepest portion of the muscle, which obviously corresponds 
-with the ulnar origin of the pronator teres in man, is slightly 
separated from the rest by the branch of the median to the 
flexor digitorum sublimis^ (see Fig. 18). 

Flexor digitorum sublimis (Fig. 18, Fl. d, s.) forms the middle 
sector. It arises from the inner part of the ulnar condyle of the 
humerus, covering the preceding sector at this part It soon lies 
on the same plane with it, occupies the middle of the forearm 
and receives an accession in the form of a portion arising by a 
flat tendon from the middle of the ulna (Fig. 19, Fl, d. s.) on 
the radial side of the flexor carpi tUnaris, The fibres of this 
portion join the under surface of the ulnar side of the general 

> I apply the tenn 'hand' to the distal segment of the fore limb simply 
for the oonvenience of distinguishing it from the distal segment of the hmd 
limb. 

* In the Seine this portion, similarly related to the nerve and arising from 
the condyle beneath the rest of the muscle, is quite separate in its whole length, 
formiiig a pronator intennediut, and, serially, corresponding precisely with the 
popliteut in ourselves. 



38 PROFESSOR HUMPHRT. 

mass. It terminates over the carpus in a broad tendon, Tvhicfa 
is joined by a portion of the deep stratum, and divides into 
four tendons to the four^ terminal phalanges. The disposition 
of the tendons corresponds almost precisely with that of its 
homologue in the hind limb (p. 26). Each subdivides into 
three, two lateral and a middle, the two lateral pass to the 
sides of the approximated ends of the metacarpal and the 
proximal phalanx, while the middle portion passes on to the 
second or terminal phalanx. In digit iv., which has three 
phalanges^ the middle portion again subdivides into two, of 
which the deeper is attached to the base of the second phalanx^ 
while the superficial runs on to the terminal phalanx. 

Flexor carpi ulnaria (Fig, 18, Fl. c, u.) arises from the inner 
condyle in conjunction with* the ^ea?or digitorum, and is inserted 
along the lower two-thirds of the ulna and into the proximal 
ulnar carpal bone; and some of its fibres are continued into 
the abductor minimi digiti which extends along the side of the 
metacarpus to the ulnar side of digit v. 

The DEEP STRATUM arises from the plantar surface of the 
ulna commencing a little below the elbow-joint, also from the 
cartilages forming the middle or radial part of the carpus, but 
not those quite on the radial side. The fibres pass partly into 
the under surface of the radial part of the jlexor svblimis digi- 
torwm aud, partl}% to the metacarpal and to the distal carpal 
bone on the radial side, also to the ulnar edge of the radius. 

More accurately described the arrangement of the compo- 
nents of this stratum is as follows in three paits — Pronator 
radii quadratics, Pronator marifdia, and Flexor profundus digi- ^ 
tarum. 

1. Pronator radii quadratua (Fig. 18, Pr, q.) arises from 
the upper part of the palmar surface of the ulna near the radial 
edge, and is inserted along the ulnar edge of the radius, its 
uppermost fibres blending at their insertion with the deepesit 
fibres of the superficial layer (pronator teres). The upper fibres 
have some obliquity downwards from the ulna to the radius, 
the lower fibres are more transverse, and rather behind the 
upper. 

^ Of the four digits in this limb digit it. only has three phalangw, the 
remaing digits ii. m. and v. having each but two plmlanges^ 



THE CBTPTOBRAKCHUS JAPONICUS. 89 

The position of the muscle on the same level with the next portion 
of the deep layer indicates, as does the passage of the large branch of 
the median behind it, that it is a segment from the deep stratum 
rather than a third and hinder stratum which one might at first be 
disposed to regard it, and which view the position of it^ correspond- 
ent in the hind limb rather favours. By the connection of the 
muscle with the lower part of the pronator teres a continuity between 
the two muscles and between the two strata is established above. It is 
not so large, does not descend so low as its correspondent in the hind 
limb, and it is not placed behind the plane of the deep flexor stra- 
tum as the pronator tibias is in that limb. The difference in the last 
particular is caused by the greater size of the pronator tiJbiai as well 
of the deep flexor stratum in the hind limb, so that the one comes 
to overlie the other. 

2. The largest portion of the deep stratum, which I will 
call 'pronator manHs* (Figs. 18 and 19, Pr. m.), arises from the 
radial side of the palmar surface of the ulna, beneath the pre- 
ceding, and from the middle carpal cartilages. Its superficial 
fibres run into the radial part of the under surface of the flexor 
sublimiSj just as the fibres from the ulnar origin of that 
muscle run into the ulnar part of its under surface. Its deeper 
fibres are inserted into the distal radial carpal bone and the 
base of the metacarpal of ii. and IIL 

3. The flexor profundus digitorum (Figs. 18 and 19) is a 
flat band arising from the upper part of the palmar surface of 
the ulna between the pronator quadratus and the fleiror carpi 
ulnaria. Like the corresponding band in the hind limb, it is 
completely segmented from the other muscles, and descends 
vertically. It passes deep, is connected with the distal carpal 
cartilage near the base of digit iv. detaching some fibres to the 
cartilage, and acquiring some from it ; indeed the cartilage pro- 
jects up through the muscle. The muscle now spreads out 
£an-like, and sends a division to each of the four digits. Each 
division is inserted chiefly into the base of the proximal phalanx, . 
and a lateral detachment passes to one or both sides of the 
metacarpal. The division which passes to digit ii. encounters 
another distal carpal cartilage, which also projects up through 
it, some of the muscular fibres passing to the cartilage and some 
•passing from it. This trumpet-like muscle does not, as in the 
hinder limb to so great an extent, curl beneath the level of 
the pronator portion of the stratum (the pronator manils)^ 



40 PROFESSOR HUMPHRY. 

because that portion is in the fore limb smaller, not arising 
from the ulnar part of the carpus; and the trumpet- like part of 
the flexor profundus digitorum is more on the same level with 
it, and lies on its ulnar and distal sides. Still the portion to 
digits II. and ill. does, as in the hind limb, curl beneath the 
superficial fibres of the pronator manHs which pass into the 
under surface of the flexor eublimis going to those digits. 

The short muscles resemble those in the hind limb (p. 28). 

Carpo-metdcarpaies pass from the distal row of the carpals, 
each to the sides and middle part of its metacarpal. The 
middle are shorter than the lateral fibres, to give space for the 

Metacarpo-phalangei which pass from the middle of the 
palmar surface of the metacarpals to the bases of the proximal 
phalanges. 

Phalangeus is present only in digit iv. passing from the 
proximal to the second phalanx. 

Dorsal Muscles of Forearm and Foot. 

On the dorsal aspect of the forearm and hand the muscles 
form a ' supinato-extensor' mass, corresponding, serially, with 
the 'supinato-extensor^ mass in the leg and foot, and, aotago- 
nistically, with the 'pronato-flexor' mass in the 1^ and foot, 
and in the forearm and hand. Like those, they are partially 
divisible into a superficial and a deep stratum. 

The SUPERFICIAL STRATUM arising from the dorsal surface of 
the radial condyle and the adjacent part of the humerus, prepon- 
derates over the deep stratum more than is the case on the 
palmar aspect, It is in three sectors, a ' radial,' an ' ulnar,' and 
an 'intermediate.' 

The radial sector representing the supinator longus and 
brevis and the extensores carpi radiales is the largest. It arises 
from nearly the whole of the part of the humerus mentioned, 
and is inserted into the whole of the dorsal surface of the radius. 
A portion of its fore part extends over the wrist-joint and car- 
pus, and is inserted into the ulnar side of the base of met. ll. 
representing the extensor carpi radialis longxor^ (Fig. 17, ExL 

' In Menobraneh it runs on to the radial carpal bone, but not to the meta- 
earpoB. 



THE CRTPTOBRANCHUS JAPONICUS. 41 

c. r.). The upper and deep part, which might be regarded aJs ap- 
pertaining to the deep stratum, represents the supinator Irevis 
and the part lying between these two, and inserted into the 
radius nearer the wrist, represents the supinator radii longus 

(Fig. 17). 

The middle sector (extensor digitorumt suhlimis) arises from 
the humerus, superficially to the radial sector. Passing down 
the forearm and over the carpus, and receiving upon its under 
surface fibres from the extensor digitorum brevis, it divides to the 
four digits, passing to the terminal phalanges \ 

The ulnar sector (extensor carpi vlnaris) arises on the ulnar 
side of the other sectors, is inserted into nearly the whole of the 
shaft of the ulna, and a portion runs on to the ulnar side of the 
carpus. 

The greater proportion of the fibres therefore pass from the 
humerus to the ulna instead of, as in most higher animals, from the 
ulna to the carpus. A similar deviation from the disposition in 
higher animals also is presented to a still greater extent by the corre- 
sponding stratum in the hind limb, the fibres of which pass largely 
from the femur to the tibia and fibula^ instead of as in Man all 
passing from the leg to the foot. 

The DEEP STRATUM is absent from the upper region of the 
forearm, except the part of it blended with and forming the 
deeper portion of the radial sector just described as represent- 
ing the supinator brevis. 

The uppermost part of the deep stratum, with that excep- 
tion, arises from the dorsal surface of the lower end of the ulna^ 
and crossing over the back of the radius and the carpus and. 
the extensor carpi radialis, is inserted into the radial side of 
the base of met. ii. It represents the extensor poll, primus. 
It may be best called ' Supinator manHs* (Fig. 17, Sup. m). 

This is a better term than that of 'rotator earrpV which I applied 
to the same muscle {Journal of AmU, iv. 48). It is one of the most 
constant muscles in the forearm, is the serial homologae of the muscle 
which I have called supinator pedis (p. 29), and, like it, is in the same 
plane with and in a higher level than the extensor brevis and often 
connected with it. 

^ In Menobranch and in Sanrians it stops, like its homologae in the hind 
Hmb of those animals and of Ai, at the metacarpns, being inserted there in three 
portions ; and smaU mnsoles arising from the metacarpus, close to its insertions, 
fonstitatc the only extensors of the digits, and pass to the terminal phalanges.' 



32 PBOFESSOR HUMPHRT. 

frontier lines into other districts, and sometimes paas from an 
extensor to a flexor office. 

On the Palbcar Aspect of the Girdle and Arbl 

The pectoral (Fig. 6, P.). is derived ahnost entirely from 
the superficial stratum of the obliquo-rectus of the abdoioen, 

and is continuous with it. Where the fibres diverge fit>m 
the trunk-muscle and take their independent course over the 
coracoid and to the humerus, they lose the tendinous inscrip- 
tions, ie. in the last four inches of their course. A few fibres 
are derived from the sternum; and as the muscle passes over 
the hinder part of the coracoid it acquires some addition from 
muscular fibres which arise from the coracoid. It is inserted 
into the radial edge of the radial tubercle near the upper end of 
the humerus, none of the fibres extending beyond this tubercle. 
Its anterior edge is continuous with fascial tissue superficial to 
the precoracoid and extending over the neck\ 

The coracoid and precoracoid cartilages in this animal are 
very large and form a considerable part of the glenoid cup : and 
the muscles arising from them are numerous, almost surround 
the head of humerus, and are difficult of interpretation. They 
arise ehiefly fix)m the outer' surface or the edge, and are as 
follows. 

1. A broad thin muscle, arising from the outer surface of 
the sternal or epicoracoid edge of the coracoid superficial to the 
biceps. It crosses the muscular fibres of the biceps superficially 
and transversely and converges to be inserted into the summit 
of the upper part of the radial tubercle of the humerus, just 
above the pectoral. It may be called eptcoraco-humeral (Fig. 
17)*. Some of its superficial fibres are blended with those 
of the under surface of the pectoral, and it is not improbably 
the representative of the pectoraUs minor of manmials. 

^ This IB the fasoial iiiTestment already spoken of (p. 2) as eoTering Uie 
body. Schmidt, Goddard, and van der Hoeyen, speak of uie conneetion of the 
pectoral with the stemnm and coracoid, bat do not mention its relatioii to the 
external oblique, or to the fascial tissue. 

* It oonesponds, I think, with that described nndw this naina in the 
Echidna, by Mivart. Tnmt. Linn, Soe, zxy. 8S8. 



I 

i 



THE CRTPTOBRANCHUS JAPOKICUS. S3 

2. Precoraeo-brachicU (Rg. 17, P. c. 6.), arises firom the whole 
of the outer surface of the precoracoid cartilage, with the excep- 
tion of the margined part. It lies in the same plane with the 
epiooraco-hnmerai, indeed is almost continuous with it, and is 
inserted, in close connection with it and with the pectoral, into 
the radial side of the uppermost part of the radial tubercle of 
the humerus *. 

3. Coraco-brachialis Uyiigus (Figs. 17 and 18, C. h. I.) is the 
largest of the muscles arising from the coracoid It arises from 
the hinder edge of the coracoid and divides into two portions. 
Of these, the larger and inner or lower division is inserted into 
the ulnar edge of the humerus for a quarter of an inch above 
the internal condyle : the other division, being nearly as large, 
is partly inserted into the side of the long tendon of the biceps, 
while a bundle of its fibres is continued on over the elbow, 

and is inserted into the ulna near the joint 

« 

This last-described division must represent the short or coracoid 
origin of the biceps in Man. There is no trace of it in Menobranch 
Axolotl or Newt. The muscle in them though large is confined to 
the humerus in its insertion. 

4. Coraco-hrachialis hrevis (Fig. 17, C. h. br.) arises from 
the coracoid close to the shoulder-joint, between the preceding 
muscle and the joint, and also from the external surface of the 
hinder part of the coracoid near the joint. It passes beneath 
the biceps to the ulnar side of the humerus near the shoulder- 
joint, and to the base of the ulnar side of the radial tubercle of 
the humerus beneath the cordco-brachiaiis auperficialia. It cor- 
responds with the ordinary mammalian coraco-brachial. 

The median nerve passes between these last two ; while the ulnar 
vessel, and in animals where it is present the ulnar nerve also, con- 
tinues its course behind both. 

^ This nrasele is eaUed tubclavius by Mivart, in his desoriptioD of Menopoma 
and Menobranch. Proc. Zool. Soc. 1869, pp. 265 and 460, thongh he regards it 
as the lame as that named by him epicoraeo-humeral in ttie Echidna. The re- 
lations of the muscle to the mammalian tuhclavius do not appear to be suffi- 
ciently clear to induce me to foUow in the application of that name to it ; uid it 
arises from the precoracoid rather than from the epiooracold part of the girdle. 
In Kenobranoh it is inserted into the summit of the radial tubercle, and its 
imder surface is blended with the supra-scapular which makes its appearance in 
that animal though quite absent in Cryptobraneh. 

YOL. VL 3 



> 

u 



Si PROFESSOR HUMPHRY. 

5. Goraco-brachialis quartus (subsoapularts) arises from the 
margin of the coracoid and also of the scapula forming the 
inner edge of the glenoid cup, extending from the origin of the 
coraco-brachiah's breuts to and over that of the triceps and also 
a very short distance upon the adjacent inner surface of the 
scapula. It separates the origin of the coraco-oUcranaiid from 
the capsule and from the origin of the triceps. The latter 
expands somewhat into the capsule^ and so separates this muscle 
behind from the capsule. With that exception it is in imme- 
diate contact with the capsule which is thin under it. It is 
inserted into the upper part of the humerus behind the pre- 
ceding, that is, just behind the ulnar edge between the coraca- 
brachialis brevis and the inner or third origin of the tricepa 
Indeed some of its fibres are continous with the former of those 
muscles in front, and with the latter behind. 

This mnscle spreading itself to a greater extent upon the internal 
surface of the scapula in some animals becomes the subscapular. It 
is called subscapular by Mivart in Meaopoma and Menobranch. In 
the latter animal its exteuiiion upon the inner surface of the scapnla 
is rather more decided than in Cryptobranch. 

6. CoracO'OlecraTialis arises, by a tendinous band, from the 
edge of the coracoid, near the joint, internal to the middle of 
the preceding muscle, which separates it from the joint It 
soon becomes muscular, descends the ulnar side of the arm dor- 
sally, joins the triceps and is inserted with it into the inner side 
of the olecranon. 

This coracoid accession to the triceps is common in Reptiles as 
well as in Urodelans. It results fi'om the large relative size of the corar- 
coid, and reminds us of the accession which tlie qtuidriceps extensor 
cruris in these animuls receives in the form of what I have called in- 
ternal recttis from the inner side of the front of the hip-joint. Only, 
in consequence of the difference in the rotation of the two limbs, the 
extensor of the forearm being directed backwards, gains its accession 
fix)m the back of the shoulder-girdle, while the extensor of the leg being 
turned forwards gains its accession fi-om the front of the pelvic girdle. 

The corctco-olecrancUis is also an example of the deviation of a 
member of the flexor ^ries to an extensor function : just as the 
biceps flexor cntris is an instance of the deviation of a member of the 
extensor series to the flexor function, the office in each instance being 
determined by the position and attachment. 

7. Coraco-radialis, or biceps (Fig. 17, -B.), arises from the 



THE CRYPTOBRANCHUB JAPONICUa 83 

external surface of the coraooid, between the epiooracobrachial 
(pecL minor) and the short coraoo-brachial, as a fan-shaped 
musdoy the fibres of which pass across the short ooraco-brachial 
and soon converge into a long tendon, which runs down beneath 
the pectoral Having passed the pectoral it receives the fibres 
of the long coraco-brachial, passes over the elbow-joint, and is 
inserted into the palmar surface of the upper end of the radius 
close to the joint. It is supplied by the nerve which perfo-. 
rates the scapula and which supplies also the superficial coraco- 
brachial 

I find the biceps corresponding with the above aud distinct from 
the hrac/iialia anlicus in Menobraneb, Newt, aud Axolotl. In the 
Newt the long thin tendon is more closely related to the braehiaUa 
aniicus, but does not seem to derive muscular fibres from it. As 
above mentioned, it does not in any of these animals derive fibres 
from -the ooraco-brctchiaUs longtts. 

The Brachialis anttcus arises from the radial side of the 
radial tubercle of the humerus, close, but on the opposite side 
of the tubercle, to the insertion of the pectoral, aud beneath the 
insertion of the dorscUis scapulce^. It passes along the shaft of 
the humerus, deriving no fibres from it, passes over the elbow 
superficial to the biceps tendon, aud is inserted about equally 
into the radius and ulna Its foremost fibres are inserted into 
the ulnar side of the radius at a short distance from the joints 
nearly in the situation corresponding with the middle of the 
insertion of the supinator longus in ourselves. Its hinder half, 
passing between this radial insertion and the insertion of the 
bicepSy is attached to the adjacent palmar surface of the ulna 



On the Dorsal Aspect of the Girdle and Ahm. 

The trapezius already described, (p. 30). 

Latieeimue dorsi arises from the fascial tissue over the dor* 
sal muscle, extending, forwards, nearly to the head, overlying the 
hinder part of the trapezius and, backwards, half way to the 
pelvis. It is a thin muscle and its edge is not very defined in 

> I may obaerre that neither in this nor in the hind limh is the snrface for 
attaefament of nrascles increased by intermuBcnlar septa extending from the 
aides of the hnmems and femoi; 

S—2 



36 PROFESSOR HUMPHRY. 

front or behind. The fibres converge and are blended with the 
upper part of the scapular origin of the triceps. Through the 
medium of it some are oonnected with the hinder mai^in of the 
scapula near the glenoid cavity ; but it has no direct attach- 
ment to the scapula^ and we cannot trace any of its fibres to the 
humerus. 

In these animals, in Keptiles and Birds, the lot, d, overlaps the 
trapeziua; the reverse being the case in mammab. 

Dorsalts scapulas (Fig. 17, D, a) arises firom the outer ex- 
panded surface of the supra-scapula. It is a long trumpet- 
shaped muscle, and is inserted into the radial side of the radial 
tubercle of the humerus beneath the insertion of the pre- 
coraco-brackial. 

This is in Cryptobranch the only muscle, with the exception of 
the few fibres of the subscapularia above mentioned, passmg from the 
scapula to the humerus. In Menobranch there is in addition a 
muscle, blended with the deeper part of the precoraco-brachial, which 
represents the gupra-seapular. The daraalis seapuicB must represent^ 
in the main, the in/rO'tpiTicUvs. Its superficial part, probably, be- 
comes in other animals developed so as to contribute to the scapular 
part of the ddioidy and its hinder part is segmented as the teres 
minor. The damculofr part of the deltoid corresponds probably, to some 
extent, with the superficial fibres of the precorctco^irackial. Thus the 
clavicular and scapular parts of the deltoid come into relation with 
the trapezme^ which is, in Urodelans, inserted into both precoracoid 
and scapula. They overlie the eupror and infrar-epinaMie and teres 
nwnor muscles, and have insertion corresponding with that of the pre^ 
corcicihbrcuihud and darscUis ecapulce. The two parts are not uncom- 
monly separate. 

The few fibres of the stibscapidaris constitute the only appearance 
of muscle upon the whole of the large extent of the concave under 
surface of the coracoids and scapula, so that the surface is free to play 
upon the convex outer surfiioe of the ventral muscle on which it lies. 

Triceps arises by a tendon from the posterior edge of the 
scapula, immediately behind the joint and in close connection 
with the capsule, which indeed it strengthens by expanding 
upon it. It soon becomes muscular and is joined by the 
latissimus dorsi. A second head arises from the radial side of 
the humerus at the base of the tubercle, and a third from the 
hinder and ulnar aspect of the humerus. These all unite; and 
the muscle is inserted into the olecranon^ being joined by the 
coraco-olecrnnulis. 



the cbtptobbanchus japonicus. s7 

Palmab Muscles of Fobeark and Hand\ 

On the palmar aspect of the forearm the muscles form, as on 
the plantar aspect of the leg, a pronato-flexor mass, which is 
partially divided into two strata. 

The SUPERFICIAL STKATUM (Fig. 18) is in three sectors. A 
radial sector representing the Pronator teres and the Fleaior 
caijn radiaUsy lai^ and thick, passes from the anterior or pal- 
mar part of the ulnar condyle, commencing close to the joint 
and extending a little way up the humerus above the condyle, 
crosses the forearm obliquely, and is inserted into an oblique 
line on the radius^ which passes across its palmar surfisM^e from 
the ulnar to the radial edge, and into the radial edge as &r as 
the wrist, also into the radial side of the two radial cartilages 
of the wrist The upper edge of its insertion is immediately 
below that of the hrachidlis anticus. The deepest portion of 
the muscle, arising from the condyle close to the joint and pass- 
ing to the ulnar edge of the radius, that is, to the upper part 
of the oblique line just mentioned, is in close contact with the 
pronator quadrcOua and at its insertion is blended with it, but 
is separated from it nearer the origin by the nerve (ubiar nerve) 
passing from the median nerve to the ulnar side of the forearm. 
This deepest portion of the muscle, which obviously corresponds 
with the ulnar origin of the pronator teres in man, is slightly 
separated from the rest by the branch of the median to the 
flexor digttorum sublimis* (see Fig. 18). 

Flexor digitorum sMimis (Fig. 18, Fl d. 8.) forms the middle 
sector. It arises from the inner part of the ulnar condyle of the 
humerus, covering the preceding sector at this part It soon lies 
on the same plane with it, occupies the middle of the forearm 
and receives an accession in the form of a portion arising by a 
flat tendon from the middle of the ulna (Fig. 19, FL d. s.) on 
the radial side of the flexor carpi vlnaris. The fibres of this 
portion join the under surface of the ulnar side of the general 

> I apply the tenn < hand ' to the distal segment of the fore limb simply 
lor the oonyenienee of distinguishing it from the distal segment of the bmd 
limb. 

* In the Sdnc this portion, similarly related to the nerve and arising from 
the eondyle beneath the rest of the muscle, is quite separate in its whole length, 
lonning a frunator inlennecltiw, and, serially, corresponding precisely with the 
popliteuM in ourseWes. 



38 PROl^ESSOR HUMPHBT^ 

mass. It terminates over the carpus in a broad tendon, which 
is joined by a portion of the deep stratum, and divides into 
four tendons to the four^ terminal phalanges. The disposition 
of the tendons corresponds almost precisely with that of its 
homologue in the hind limb (p. 26). Each subdivides into 
three, two lateral and a middle, the two lateral pass to the 
sides of the approximated ends of the metacarpal and the 
proximal phalanx, while the middle portion passes on to the 
second or teiminal phalanx. In digit iv., which has three 
phalanges, the middle portion again subdivides into two, of 
which the deeper is attached to the base of the second phalanx^ 
while the superficial runs on to the terminal phalanx. 

Flexor carpi ulnaris (Fig. 18, FL c. w.) arises from the inner 
condyle in conjunction with" the ^e^ror digitoruvi, and is inserted 
along the lower two-thirds of the ulna and into the proximal 
ulnar carpal bone; and some of its fibres are continued into 
the abductor minimi digiti which extends along the side of the 
metacarpus to the ulnar side of digit v. 

The DEEP STRATUM arises from the plantar surface of the 
ulna commencing a little below the elbow-joint, also from the 
cartilages forming the middle or radial part of the carpus, but 
not those quite on the radial side. The fibres pass partly into 
the under surface of the radial part of the fleocor sublimie digi- 
torum and, partly, to the metacarpal and to the distal carpal 
bone on the radial side, also to the ulnar edge of the radius. 

More accurately described the arrangement of the compo-; 
nents of this stratum is as follows in three pai-ts — Pronator 
radii quadrattis, Pronator manHs, and Flexor profundus digi- ^ 
torum. 

1. Pronator radii quadratue (Fig. 18, Pr, q.) arises from 
the upper part of the palmar surface of the ulna near the radial 
edge, and is inserted along the ulnar edge of the radius, its 
uppermost fibres blending at their insertion with the deepest 
fibres of the superficial layer {pronator teres). The upper fibres 
have some obliquity downwards from the ulna to the radius, 
the lower fibres are more transverse, and rather behind the 
upper. 

^ Of the four digits in this limb digit iv. only has three |khftlangM, the 
xemaing digits xi. ui. and t. having eeuUi but two phalanges^ 



r 



THE CKTPTOBRANCHUS JAPOKICUa S9 

The position of the muscle on the same level with the next portion 
of the deep layer indicates, as does the passage of the large branch of 
the median behind it, that it is a segment from the deep stratum 
rather than a third and hinder stratum which one might at first be 
diAposed to regard it, and which view the position of itn correspond- 
ent in the hind limb rather favours. By the counection of the 
muscle with the lower part of the pronator teres a continuity between 
the two muscles and between the two strata is established above. It is 
not so laige, does not descend so low as its correspondent in the hind 
limb, and it is not placed behind the plane of the deep flexor stra- 
L turn as the pronator tibias is in that limb. The difference in the last 

particular is caused by the greater size of the pronator tihixB as well 
of the deep flexor stratum in the hind limb, so that the one comes 
to overlie the other. 

SL The largest portion of the deep stratum, which I will 
call * pronator manUs* (Figs. 18 and 19, Pr, m.), arises from the 
radial side of the palmar surface of the ulna, beneath the pre- 
ceding, and from the middle carpal cartilages. Its superficial 
fibres run into the radial part of the under surface of the flexor 
sublitnia, just as the fibres from the ulnar origin of that 
muscle run into the ulnar part of its under surface. Its deeper 
fibres are inserted into the distal radial carpal bone and the 
base of the metacarpal of il. and IIL 

3. The flexor profundtis digitarum (Figs. 18 and 19) is a 
^ flat band arising from the upper part of the palmar surface of 

' the ulna between the pronator quadratu8 and the flsxor carpi 

tdnar%8. Like the corresponding band in the hind limb, it is 
completely segmented from the other muscles, and descends 
vertically. It passes deep, is connected with the distal carpal 
cartilage near the base of digit IV. detaching some fibres to the 
cartilage, and acquiring some from it ; indeed the cartilage pro- 
jects up through the muscle. The muscle now spreads out 
fan-like, and sends a division to each of the four digits. Each 
division is inserted chiefly into the base of the proximal phalanx, 
and a lateral detachment passes to one or both sides of the 
metacarpal. The division which passes to digit ii. encounters 
another distal carpal cartilage, which also projects up through 
it, some of the muscular fibres passing to the cartilage and some 
•paasing from it. This trumpet-like muscle does not, as in the 
^ hinder limb to so great an extent, curl beneath the level of 

the pronator portion of the stmtum (the pronator manHs), 



40 PBOFESSOR HUMPHRY. 

because that portion is in the fore limb smaller, not arising 
from the ulnar part of the carpus; and the trumpet- like part of 
the flexor profwndua digitorum is more on the same level with 
it» and lies on its ulnar and distal sides. Still the portion to 
digits II. and III. does, as in the hind limb, curl beneath the 
superficial fibres of the pronator manila which pass into the 
under surface of the flexor sublimia going to those digits. 

The short muscles resemble those in the hind limb (p. 28). 

Carpo-metacarpaies pass from the distal row of the carpals, 
each to the sides and middle part of its metacarpal. The 
middle are shorter than the lateral fibres, to give space for the 

Metacarpo'phalangei which pass from the middle of the 
palmar surface of the metacarpals to the bases of the proximal 
phalanges. 

Phalangeus is present only in digit lY. passing from the 
proximal to the second phalanx* 

Dorsal Muscles of Forearm and Foot. 

On the dorsal aspect of the forearm and hand the muscles 
form a ' supinato-extensor' mass, corresponding, serially, with 
the 'supinato-extensor^ mass in the leg and foot, and, antago- 
nistically, with the 'pronato-flexor' mass in the 1^ and foot, 
and in the forearm and hand. Like those, they are partially 
divisible into a superficial and a deep stratum. 

The SUPERFICIAL STRATUM arising from the dorsal surface of 
the radial condyle and the adjacent part of the humerus, prepon- 
derates over the deep stratum more than is the case on the 
palmar aspect, It is in three sectors, a ' radial,' an ' ulnar,' and 
an 'intermediate.' 

The radial sector representing the supinator longus and 
brevie and the exteneorea carpi radiates is the largest. It arises 
fix)m nearly the whole of the part of the humerus mentioned, 
and is inserted into the whole of the dorsal surface of the radius. 
A portion of its fore part extends over the wrist-joint and car- 
pus, and is inserted into the ulnar side of the base of met. n. 
representing the extensor carpi radialia longior^ (Fig. 17, -Ect 

1 In Menobraneh it runB on to the radial earpal bone, bat not to the meta- 
earpoB. 



THE CBTPTOBRAKCHUS JAPONICUS. 41 

e. r.). The upper and deep part, which might be T^;arded as ap- 
pertaining to the deep stratum, represents the supinator brevis 
and ihe part lying between these two, and inserted into the 
radius nearer the wrist, represents the supinator radii longus 
(Fig. 17). 

The middle sector {extensor digitorwm sublimis) arises from 
the humerus, superficially to the radial sector. Passing down 
the forearm and over the carpus, and receiving upon its under 
surface fibres from the extensor digitorum hrevis, it divides to the 
four digits, passing to the terminal phalanges^ 

The ulnar sector (extensor carpi ulnaris) arises on the ulnar 
side of the other sectors, is inserted into nearly the whole of the 
shaft of the ulna, and a portion runs on to the ulnar side of the 
carpus. 

The greater proportion of the fibres therefore pan from the 
humerus to the ulna instead of, as in most higher animalci, from the 
ulna to the carpus. A similar deviation from the disposition in 
higher animals also is presented to a still greater extent by the corre- 
sponding stratum in the hind limb, the fibres of which pass largely 
from the femur to the tibia and fibula, instead of as in Man all 
passing from the leg to the toot. 

The BEEP STRATUM is absent from the upper region of the 
forearm, except the part of it blended with and forming the 
deeper portion of the radial sector just described as represent- 
ing the supinator brevis. 

The uppermost part of the deep stratum, with that excep- 
tion, arises from the dorsal surface of the lower end of the ulna, 
and crossing over the back of the radius and the carpus and. 
the extensor carpi radialis, is inserted into the radial side of 
the base of met. il« It represents the extensor poll, primus. 
It may be best called ' Supinator manils* (Fig. 17, Sup, m ). 

This is a better term than that of Wotator carpi* which I applied 
to the same muscle {Journal of Anat, iv. 48). It is one of the most 
constant muscles in the forearm, is the serial homologue of the muscle 
which I have called supinator pedis (p. 29), and, like it, is in the same 
plane with and in a higher level than the extensor brevis and often 
connected with it. 

I In Menobranoh and in Saorians it stops, like its homologne in the hind 
Hmb of those animals and of Ai, at the metacarpns, being inserted there in three 
portions ; and smaU mnscles arising from the metacarpus, elose to its insertions, 
eonstttate the only extensors of the digits, and pass to the terminal phalanget.i 



42 PROFESSOR HUICPHBT. 

The extensor digitorum hrems or profundus is a broad thin 
muscle arising from the proximal row of carpal bones, and join- 
ing the under surface of the extensor digitorum sublimis just 
above its division. It is connected with the preceding by a 
slip from the lower end of the ulna, which passes with it to the 
under sur^GLce of the ulnar part of the extensor digitorum sublimis, 

Interossei are directed between the metacarpals descending 
rather lower. on the ulnar than on the radial sides of the several 
bones. 

The similarity in the disposition of the muscles in the middle and 
distal negments of the fore and hind limb is very closely in accordance 
with the Bimilarity in the general structure and fanctions of the limbs* 
In each they coustitute a 'pronato-flexor' mass upon the palmar or 
plantar aspect, and a 'supinato- extensor' maBsu])on the doi'sal aspect; 
and the segmentation of these masses is much alike in each. 

Esspecially in this so in the case of the supinato-extensor mass, 
of which the division into two strata is equally distinct in each limb, 
and the superficial strcUum arising from the condyle of the femur or 
humerus on the * tibial* or 'radial* side, is in three parts, of which the 
middle passes to the digits, while the laterals are attached to the 
bones of the leg and forearm, and run on to the tarsus and carpus. 
The deep stratum in each limb is composed of a supinator of the foot 
or hand continuous with a short extensor of the digits. 

In the PBONATO-FLEXOR MASS there is rather more difference in 
the two limbs, which is referable chiefly to the fact that segmentation 
is somewhat more advanced in the fore limb. In it the superficial 
strahim is more scfiarate from the deep, though the two are still 
blended above and below; and the superficial stratum presents the 
three sectors — *radial,* 'ulnar' and * inter mediate' — which is not the 
case in the hind limb, where the muscular force is concentrated 
' chiefly upon the flexion of the distal parts, and is less expended on 
the pronation of the tibia. This stratum moreover in the hind limb 
receives an accession from the tail and from the pelvis. With that 
exception the origin in the two limbs is alike — from the fibular con- 
dyle of the femur, and the fibula in the one, and from the ulnar 
condyle of the humerus, and the ulna in the other. The de^ strata 
in the two correspond with an exactness which is scarcely modified 
by the difference in the number of the digits, each consiijting, after 
the pattern of the dorsal antagonistic stratum, of a pronator of the 
foot or hand continuous with a deep flexor of the digits. 

There ib, however, this difference between the antagonistic deep 
strata, viz. that a distinct vertical band or strip is in both limbs 
segmented from the whole length of the pronator portion, and joined 
more or less completely to the short flexors constituting that which 
I have called the deep flexor of the digits; whereas, in the dorsal 
aspect, the short or deep extensor is connected only with the lower 
edge of Uie supinator. 



THE CBTPTOBBANCHU8 JAPONICUS. 4S 



MUSCLES OF THE HEAD. (Figs. 5 and 20.) 

I have already (p. 30) de8cri}>ed the subciUaneuB colli. 
Beneath it is the depressor mandihulcB, presently to he noticed. 
Next lies the constrictor faucium, which arises from the fascial 
tissue between the depressor mandibiUoB and the trapezius^ 
Narrow at first, it radiates out beneath the neck and the hinder 
part of the broad interval between the mandibular rami and 
meets its fellow of the opposite side at the middle line. 

The Mylo-hyoid arises from the hinder extremity of the 
comu of the hyoid and the ligament^ which connects the 
hyoid comu with the suspensory apparatus of the jaw, as well 
as from the suspensory (glenoid) cartilage where it projects 
behind between the mastoid and the pterygoid bones, but not 
from the angle of the jaw itself, being separated from it by the 
insertion of the digastric. It passes forwards and radiates 
on the deeper surface of the subcutaneous muscle, and in con* 
nection with it is inserted into the inner part of the ramus of 
the lower jaw. 

Oenio-branchial, a long muscle, passes from the hinder part 
of the posterior branchial cartilage to the hinder surface of the 
lower jaw, at a little distance from the middle line. 

Branchio-hyoid, a thick muscle, arises from the hinder part 
of the posterior branchial cartilage, its origin being by tendi- 
nous fibres from the deeper surface of the cartilage and by 
numerous muscular fibres which almost entirely enclose the 
cartilage; it arises also from the hinder part of the anterior 
branchial cartilage, and runs forwards to be inserted into the 
approximated parts of the basis and comu of the hyoid. 

Intermandibular consists of fibres passing transversely across 
the middle line from one side of the lower jaw to the other 
beneath it. 

Temporal is a large muscle arising by a broad tendon from 
the frontal and parietal bones, near the middle line of the 
skull, and by a thick portion running backwards in the groove 

^ In the bftok of that ligunent there is a distinet though small piece of earti- 
lage (stylo-byal) between the extremity of the coma (cerato-hyal) of the hyoid 
«ttd the mastoid part of the enspenftorial cartilage. Hyrtl fotmd no cartilage 
in this ligament, though he menti<mB one as present in Menopoma. 



44 PROFESSOR HUMPHRT. 

internal to the suspensory apparatus (the squamous bone and 
the ridge in the parietal upon which it abuts) to the dorsal 
spines of the three foremost vertebraB, from which it arises by 
means of a strong tendon. It forms a thick mass, bounding 
the orbit behind, and filling up the wide pterygoid fossa on the 
side of the skull, but not deriving many fibres from the bones 
there. The fibres converge to be inserted into the upper surface 
of the lower jaw, and particularly the inner or * surangulax' part 
of it, a little in front of the articulation with skulL Some of 
the fibres arising from the pterygoid probably represent the 
external pterygoid muscle. 

Afasseter, a thick muscle, arises tendinous from along the 
upper surface of the bpne ('squamous/ as it appears to me), 
forming the upper part of the suspensorium, and descends to 
be inserted into the outer surface of the lower jaw, for an inch 
in front of the joint. — A considerable mass of muscle arises 
from the anterior surface of the suspensorium and is inserted 
into the upper surface of the jaw between the temporal and 
the mastoid. It is partly separated from the masseter by 
areolar tissue, in which branches of the third division of the 
fifth pass outwards to the side of the head and to the lower jaw. 
Its fibres are, however, blended to some extent with those of the 
masseter and of the temporal; and I cannot tell whether it 
should be regarded as appertaining more particularly to either, 
or to both, or as constituting a distinct pterygoid muscle. 

Digastric, a strong muscle, arises from the retiring angle 
between the suspensorium and the back of the skull behind, 
near the auricular opening, and from the edges of the squamous, 
the mastoid, the parietal and exoccipital as well as the cartilage 
which they there surround. At its origin it is between the 
origin of the masseter from the squamous bone and the inser- 
tion of the dorsal muscle of the trunk into the exoccipital 
(Fig. 20). It lies on the groove of the mastoid along the back 
of the suspensorium, and is inserted into the hinder part of the 
angle of the lower jaw, just above the depressor maxillse; but 
a distinct and strong bundle of its fibres passes on tendinous 
behind the jaw along the hinder surface of the suspensory liga* 
ment of the hyoid to the cornu of the hyoid. It is here over- 
lapped by the mylo-hyoid, or those fibres rather of it which 



THE CBTPTOB&ANCHUS JAFPmCUH. 49 

are attached to the suspensory ligament of the hjoid and the 
suspensory cartilage of the jaw. 

The close conQection of these parts — angle of jaw and its suspen- 
aoiial cartilage — with the oomu of the hyoid aud its suspensorial 
ligament, together with the insertion of the digastric into the angle of 
the jaw and the hyoid corn a, are interesting and important in reUition 
to the remarkable course of the digastric to the hyoid and to the 
jaw, in ourselves. 

Depressor Mandibulcs arises two inches behind the head 
from the fascial tissue, near the middle line, beneath the suhcu* 
taneaus colli, between it and the constrictor faucium, forms a 
band about half an inch broad, the fibres of which converge to 
be inserted tendinous into the extremity of the jaw close to the 
preceding. 

THE NERVES OF THE HEAD (Fig. 20). 

The tough areolar tissue of the animal and the numerous 
accompanying veins rendered the dissection of the nerves 
difficult. 

For an account of the nerves in the cranial cavity, I may 
refer to Figure 22 and the description of it. 

The Optic passes from the skull by a separate foramen^ 
through the orbit, to the eye. Mr Anningson, in careful dis- 
section of both orbits, could not discover the third, fourth, or 
sixth nerves in the orbit The third and fourth were however 
found in the cranial cavity very small, and are represented in 
Fig. 22. Schmidt, Goddard, and van der Hoeven state that 
the sixth is a branch from the fifth in the orbit This we did 
not find. 

The three divisions of the fifth nerve are quite distinct. 

The first, or ophthalmic division, and the second, or SUPRA- 
MAXILLARY, escape from the skull by a common hole, though 
separated by a fibrous band. 

The OPHTHALMIC is a large nerve, crosses above the optic» 
gives a suprOrorbitcU branch, which passes to the integuments 
above the eye, and which, on its way, supplies nerves {ciliary) 
to the eye. Soon after entering the orbit, the ophthalmic de- 
taches a large branch which runs beneath the eye, and at the 



a PROFESSOR HUMPHRY. 

« 

margin of the orbit joins a long branch from the third division 
of the fifth. The resultant trunk passes through an iufra»or- 
bital hole, and is the infra-orbital nerve. Many small branches 
are given off by both nerves before they unite. The ophthalmic 
also, immediately after crossing the optic, gives a branch (nasal) 

to the nose. 

The SUPRA-MAXILLARY, or second division of the fifth, runs 
along the floor of the orbit beneath the optic nerve and divides 
into branches, which pass inwards to the nose and forwards to 
the maxilla. 

The INFRA-MAXILLARY, or third division of the fifth, emerges 
through a large hole at the front of the base of the suspensorial 
projection for the jaw, between the temporal and masseter 
muscles, and gives branches to both. Its dentary brahch runs 
outwards beneath the masseter muscle, between it and the 
muscle arising from the front of the suspensorium which it 
supplies, then dips down to the lower jaw, and enters it by 
two branches through two foramina. A branch of the infra- 
maxillary runs forwards over the temporal muscle, on the outer 
side of the eye, gives off numerous branches upon the upper 
jaw, and joining a branch of the ophthalmic, as above described, 
forms the infra-orbital. A branch passes backwards beneath 
the masseter where it comes into contact with the seventh 
nerve, communicates with it, runs outwards with it, and turns 
forward along the outer side of the ramus of the lower jaw. 

The SEVENTH nerve emerges from the skull in the retiring 
angle between the suspensorium and the occiput, and divides 
into two chief branches. One of these pierces the digastric 
muscle, supplies it, the depressor mandibuke, the subcutaneus 
colli, and the constrictor faucium. The other chief branch 
passes beneath the digastric upon the suspensorium and along 
it, communicates with the branch of the fifth which has come 
beneath the masseter to this point, passes over the ramus of 
the lower jaw, and runs along beneath it supplying the mylo- 
hyoid and the subcutaneus colli, and reaching the interman- 
dibular muscle. A branch also passes along the side of the 
lower jaw in company with the branch of the third division of 
the fifth. 

The Vagus on its first appearance in the neck is large, and 



\ 



THE CRTFTOBRANGHUS JAFONICUS. 47 

immediately gives off branches to the hrandiio-hyaul muscle 
and probably to the other branchial muscles (though these were 
not made out), small branches to the digastric and depressor 
ntandtbtilcB, the constrictor Jiziunu/m, and a large branch to the 
trapezius, also a long LATERAL nerve, which runs backwards 
along the lateral septum, in company with the lateral vessels, 
continuing its course to the tail without, apparently, giving off 
any branches. Having arrived at the tail, it gives off branches 
and communicates with the other nerves. 

The branches above mentioned correspond with branches in our- 
selves finom the 7th and 9th, and with the spinal accessory. 

The office of the lateral nerve is probably to harmonize and 
prodace simultaneous action of the several segments of the lateral 
musde. It differs in the Cryptobranch from the same nerve in the 
Fi^h in that its distribution is limited to the lateral muscle of the 
tail, which indicates that the simultaneous jiowerful contraction of 
the parts of the lateral moscle in this animal are confined to or take 
place chiefly in the taiL 



THE SPINAL NERVES. 

The several spinal nerves, emerging through the neural 
foramina behind the transverse vertebral processes, cross 
above (dorsally, with regard to) the deep lateral vessels 
which traverse the holes in the roots of the transverse pro- 
cesses, and lie above the lateral septum. They supply the 
several segments of the trunk-muscles, taking their course 
behind the respective fibrous septa or iDscriptions, and in addi- 
tion send nerve-trunks to the limbs. Each nerve, immediately 
after its emergence from the vertebral foramen, above the 
lateral septum, divides into a posterior or dorsal, and an 
ANTERIOR or VKNTRAL SPINAL nerve (Fig. 21). The former 
divides into branches, which are directed upwards and outwards, 
in front and behind the articulating processes, to the dorsal 
muscles and the skin. The cutaneous branch (Fig. 21, C) runs 
in front of the articulating processes outwards with a slight in- 
clination upwards, behind the rib. and the transverse septum 
passing dorsally from it, to the skin. The muscular branches 
radiate into the surrounding dorsal muscle. The anterioe 
SPINAL or ventral nerves (v) pass through the lateral septum 



48 PROFESSOR HUMPHRT. 

80 afi to come into contact witb the subvertebrai rectus and its 
prolongations, and take their couFBe behind the several ribs 
and the transverse intermuscular septa, and supply the ventral 
muscles, the skin and the limbs. 

The first cervical nerve (suboccipital) passes through a 
foramen in the atlas direct to the foremost portion of the suh^ 
vertebrai rectus and the adjacent dorsal muscle. The second 
cervical nerve passes to the next portion of the subvertebrai 
rectus, and a branch traversing that muscle reaches the 3rd 
nerve and so enters into the formation of the brachial plexus. 

The BtiACHiAL plexus (Fig. 4) is formed by branches from 
the 3rd, 4th, 5th and 6th spinal nerves, with that just men- 
tioned from the 2nd. 

The third cervical nerve, joined by the branch from the 
second, divides into four. No. 1 joins the fourth spinal nerve. 
No. 2 passes in front of the precoracoid and is lost apparently 
in the areolar tissue and skin of that region. No. 3 traverses 
the coracoid hole and supplies the coraco-brachialis superfi- 
cicdis, the biceps^ and perhaps the coraco-brachialis breuis. No. 4 
supplies the omo-hyoid. 

Its branches correspond with some of those which in man pass 
from the upper part of the cervical plexus, and form the external 
CUTANEOUS or MUSCULO-CUTAN Eous mass; and they correspond serially 
with the OBTURATOR nerve in the hind limb. The branch to the 
omo-hyoid is to be associated with the absence of a hypoglossal nenre: 
at least we could not discover that uorve in the dissection of the 
neck or in the interior of the skull. 

The fourth spinal nerve crosses behind the origin of the ex- 
ternal oblique from the second rib, is connected with the third 
and fifth nerves by commissural trunks, and gives off scapular 
branches, which pass beneath the scapula (subscapular) also 
to the latissimus dorsi, dorsalis scapulce and precoraco-brachial 
muscles, and a cutaneous branch (circumflex) to the outer 
side of the arm. 

The nerves to the last two muscles might be designated supra- 
scapular because they supply the muscles (dorsalis scapuUs and 
preeoraeO'brachiaTj which must, iu part any rate, answer to the 
muscles (ir^frct- and suprorSpirhcUus)^ which are, in ourselves, supplied 
by the tsupra-scapular nerve. It ii interesting to observe the nerves 
in this animal taking a course, behind the scapula to supply the 






THE CRTPT0BBANCHU8 JAPONICUS. 49 

muacleB on the donnm of the scapula, corresponding with the conrBe, 
behind the ilium, which is usually taken by the gluteal nerves in 
Uieir passage to the muscles on the dorsum of the ilium. 

The same nerve supplies the triceps, and perforating the long 
and the outer heads of that muscle descends, as the MUSCULO- 
8PIBAL or RADIAL, between them and the humerus, then be- 
tween the triceps and the brachialis anticus, and then between 
the latter and the extensor muscles arising from the outer 
condyle. It supplies those muscles, and divides into two 
branches (Fig. 17), of which one perforates the extensor mass 
of muscles, becomes superficial, and runs along tho radial edge 
of the forearm and the radial side of digit IL (digit I. is missing) : 
the other branch crosses deeply, between the extensor mass and 
the radius, to the back of the forearm, where it communicates 
with a posterior interosseous branch from the median and with 
the poeterior ulnar branch of the median (the branch that is of 
the median which has descended through the triceps). It then 
descends the back of the forearm, under the extensor muscles, 
perforates the supinator mcmUs, and divides, supplying the 
apposed sides of digits II. and ill., also the apposed sides of 
digits ra. and IV. (Fig. 17). 

The co\ir8e and disposition of this nerve oorresponds closely with 
those usually presented by the radial in higher animals, even to the 
share it takes in supplying the digits. It is derived from the middle 
of the brachial plexus in connection with the circumflex and subsca- 
pulaTy and is distributed exclusively to the muscles upon the dorsal 
aspect of the limb. 

The fifth spinal nerve is connected by a commissural branch 
with the fourth, and receives the greater part of the sixth, that 
nerve merely sending off, besides, a cutaneous branch. It 
passes between the origins of the external oblique, from the 3rd 
and 4th ribs, and supplies branches to the pectoralis and coraco- 
hrachiaUs Umgvs and probably brachioMs amUcus^ though the 
branch was not traced to that muscle. High in the arm it 
sends a long nerve (posterior ulnar, or, better, inferior 
MUSCXTLO-spinal) which, tuiTiing outwards to the back of the 
arm, perforates the short or ulnar origin of the triceps and 
descends through the triceps to the radial side of the olecranon. 
This nerve (Fig. 17) then pierces the extensor carpi ulnaris, and 

vol. VI. 4 



50 PROFESSOR HUMPHRY. 

having reached its deeper surface and communicated with the 
radial and with the posterior interosseous from the median, 
runs along the dorsal surface of the ulna^ then over the carpus, 
supplies the small muscles in that situation upon the carpus 
and metacarpus, and is distributed to the ulnar side of digit v. 
and the apposed sides of digits IV. and V. 

The main trunk of the fifth spinal nerve continues its course 
as the MEDIAN on the inner side of the arm. It passes over the 
upper surface of the humeral portion of the coraco-brcLchialis 
loTiffus, between it and the coraco-brachialis hrevis, keeps be- 
neath the bicipital portion of the former muscle, and passes over, 
the elbow between the tendon of the biceps, which is inserted 
into the radius and the portion of the coraco-hrachialia loriffus 
which is inserted into the ulna. At the elbow it divides into 
two trunks, an anterior ulnar and a median, both of which 
disappear beneath the mass of flexor muscles arising from the 
inner condyle. 

The ULNAR trunk (Figs. 18 and 19) takes its course 
through the flexor mass on the forearm, between the superficial 
stratum arising from the condyle and the ulna and the deeper 
stratum arising from the ulna and supplies both strata. One 
of its nerves passes to the Jlexor dig. subl. through the pro- 
nator teres, or proximal part of the condyloid stratum, just as 
the median in man passes between the condyloid and the 
ulnar origin of that muscle. Its terminal branch travels down 
between the flexor svhl dig. and the Jlexor prof, and, partly, 
also under cover of the flexor carpi ulnaris, supplying them in 
its course to the wrist. It runs over the carpus, supplies the 
ulnar side of dig. v., dips down between the muscles to dig. v. 
and, like the external plantar nerve in the hind limb, curls 
in beneath the fl>ea>or projimdus dig., and passes across towards 
the radial side^ of the limb, between that muscle and the meto- 
carpo^hcUangei, supplying the latter. At the interval, be- 
tween digits V. and iv., it sends down a branch which supplies 
the apposed sides of those digits, and at the intervals between 
rv. and in. and between iiL and n., it communicates with the 
branches of the median which are passing to supply the 
apposed sides of iv. and iii. and III. and H. 

The MEDIAN (Fig. 19) trunk takes a still deeper course be- 



THE CRYPTOBRANCfHUS JAPONICUS. 51 

Death the pronator quadratus, supplying it, then over the front 
of the carpus beneath the flexor profundus, where it divides 
into three nerves, one supplying the muscles of the base of 
digit II. and perhaps passing on to the radial side of that digit ; 
the second supplying the apposed side of digits il. and III., and 
the third supplying the apposed sides of digits III. and rv. The 
second and third nerves last mentioned each receives a twig 
from the terminal branch of the ulnar which runs across the 
metacarpal bones and muscles. 

The ULNAB nerve always arises from the plexus iu company with the 
MEDIAN, and not uncommonly forms one trunk with it, not separating 
from it till the forearoL In the Cryptobranch the dorsal or posterior 
part of it shows a tendency to associate itself in its course with the 
radial or dorsal nerve-trunk of the limb forming the inferior musculo- 
apiraly which corresponds in mammals to the posterior branch of the 
ulnar in the fore limb, and to the external saphenous in the hind 
limb. The palmar or anterior part of the ulnar carries with it the 
muscular branches usually given off directly from the msdiak ; and 
the trunk of the median is continued on in a deep plane as the 
ANTERIOR INTEROSSEOUS to the carpus and digits. 



Nerves to the Hind Limb. 

The nerves to the hind limb are derived from four spinal 
nerves (a, b, c, d). Of these the foremost (a) sends a branch to 




(b) and runs downwards as the OBTURATOR nerve, pierces the 
pecUneua where it is taking origin from the interior of the 
pelvis, traverses the small obturatar hole in the pelvic shield, 
and supplies the adductor muscles arising from that shield. 
Whether it extends to the superficial stratum {gracilis^ &c.) 
there situated, could not be ascertained with certainty. A 
branch from it however passes backwards to join the branch of 
the sciatic, which enters the hinder {flexor cruris) part of that 
stratum. 

4 — 2 



52 PROFESSOR HUMPHRY. 

This connection with the sciatic is interesting in relation to the 
fact that the correspondent of the obturator nerve in the fore limb— 
the perforator of the coracoid or musculo-cutaneous — supplies in part 
or wholly the flexors of the forearm. 

The second nerve (b) sends a branch to unite with the third 
and fourth (c and d) in forming the sciatic and then descends, 
over the brim of the pelvis upon the Uicums muscle, as the 
ANTERIOR CRURAL nerve. It supplies the internal recttiSy the 
iliacus and the pectineua, A long branch from it (internal 
saphenous) takes a deep course between thi^ pectineus and the 
femur to the tibial side of the limb, where it emeiges between 
the gracilis and internal rectus and descends on the tibial side 
of the knee and leg. 

The deeper course down the thigh than in ourselves of the safhb- 
Kous nerve is consequent on the extension of the pectineus down- 
wards nearly to the knee (p. 20), so that the nerve has to pass under 
it to reach its wonted place of emergence. 

The third and fourth nerves (o and d) pass from the spinal 
column, the one in front and the other behind the level of the 
ilium, unite and are joined by a branch from the second nerve 
(J) to form the SCIATIC, which emerging between the ischio- 
femoral and the gluteus or pyriformisy gives oflF several branches 
to the surrounding muscles, dorsal and plantar — ^to the pyri- 
fonnis, glutei, gluteo-rectus and biceps, to the caudo-femoraZ^ 
caudo'crural and caudo-pedal, and to the hinder part of the 
flexor cruris {semitendinosus and semimembranosus. 

It gives off also the musculo-spiral or peroneal nerve 
(Fig. 10), which descends in the thigh under cover of the hiceps^ 
supplying it and passing above or in front of it at the knee, 
then nms beneath the percneus, between it and the fibula^ to 
join a large branch from the ANTERIOR tibial and supply the 
peronei and the extensor digitorum longus. It descends on the 
front of th^ leg beneath the ext, digitorum superficial to the 
extensor haUucis and on the fibular side of the extensor brevis^ 
and sends nerves to the fibular side of digit Y. to the apposed 
sides of digits IV. and v., and ends in a nerve which joins the 
branch of the anterior tibial in the interval between digits nr. 
and IV, 



THE CRYPTOBRANCHtJS JAPONICUS. 53 

The PEROKBAL or MUSCT7LO-8PIRAL neiTe answers to the radial or 
xuscuLO-SPiRAL of the fore limb, and not at all to the ulnar, though 
its name might seem to imply the oontnirj. The differenceB between 
the musculo-spirals in the two limbs, that is between the peroneal 
and the radial, are dependent chiefly or entirely upon the difference 
in the direction of the rotation in the two limbs causing a variation 
in the direction best adapted for the nerve-coarse. This is probably 
the reason that the peroneal does not traverse the whole thickness 
of the quadricepSy bnt only that part segmented from it which 
becomes the biceps. Indeed where the rotation of the limb is com- 
plete, as in Man, the nerve does not traverse even the hicept, as it 
does in these lower animals, but only gets under cover of it or keeps 
along its hinder edge. For the same reason the nerves to the qtuid- 
rieeps pass in front of the pelvis as anterior crural, while those to 
the tric^ pass behind the shoulder-girdle bound up in the radial. 

The SCIATIC nerve (Fig. 10) then divides into two, which in 
the left limb pass on the two sides of the femoro-Jlbultr, and 
are joined by a connecting trunk beyond it, so as to encircle the 
muscle. (In the right thigh the sciatic trunk passes on the 
tibial side of the femoro-fibular and divides subsequently.) 
The outer division (external popliteal) supplies the femoro- 
fibular, sends a (sural) branch to the stratum of the flexor 
mass arising from the fibular condyle, passes between the 
condyloid and fibular part of the superficial stratum, descends 
between the two strata along the fibular edge of the flexor pro- 
fundus diffitorum (Fig. 11) between it and the origin of the 
flexor sublimis from the fibula. It passes beneath the tarsus, 
curls over (deeper than) the flexor profundus, and then runs 
across the metatarsus as far as digit ii. between the flexor-pro- 
fundus and the metatarso-phalangei. In its course it supplies 
the fibular portion of the flexor sublimis, the whole of the deep 
flexor stratum (tibialis posticus, flexor profundus, flexor-halluxns 
and accessorius), the small muscles in the sole, the fibular side 
of digit v., the apposed sides of iv. and V. Between digits 
III. and IV. and between il. and iii. it joins branches from the 
POSTERiOB TIBIAL, and the combined nerves supply the apposed 
surfaces of these digits. 

The inner division (INTERKAL POPLITEAL, Fig. 10) in the 
left limb pierces the caudo-femoral close to its attachment to 
the femur (in the right limb it passes on the fibular side of this 
tendon going between it and the femoro-fibidar). It gives off 



54 FBOFESSOB HUMFEQtT. 

a long branch which descends, gives filaments to the lower part 
of the flexor cruris {semitendinosus and semimembrcmosiui), per- 
forates it, and runs down on the tibial side of the leg and foot, 
accompanying the internal saphenous towards, and probably 
upon, the tibial side of the hallux. The latter part of its 
course corresponds with that of the superficial branch of the 
radial nerve upon the radial edge of the forearm. The next 
branches from the internal popliteal, in conjunction with branches 
from the external popliteal, supply both strata of the flexor 
mass. The nerve (Figs. 10 and 11) passes deeply beneath (in 
front of) the flexor mass and the pronator tibics or popUtens 
which it supplies, gives off the anterior tibial, descends as 
POSTERIOR TIBIAL behind the tarsus and metatarsus, and divides 
into two branches, which, joined by branches firom the external 
popliteal, supply the intervals between digits I. and IL and IL 
and III.; also, probably, the deep muscles on the tibial side of 
the sole. 

The ANTERIOR TIBIAL nervo supplies the tibialia antums, 
descends over the ankle and tarsus, and divides into two nerves; 
one passes superficially to the extensor haUiids over the tarsus 
towards, and we believed to, the interval between and the ap- 
posed sides of digits I. and iL; the other passed beneath the 
extensor hallucisy supplying it, and divides into nerves to the 
apposed sides of digits n. and in. and of ill. and iv., these 
nerves being joined by the terminal branches from the peroneal. 

The similarity in the course and disposition of the external and 
INTERNAL POPLFTEAL nerves and their pkoitar branches with the course 
and diapoflition of the ulnar and median nerve and their palmar 
branches is what might have been anticipated from the similarity of 
the muscles in the corresponding parts of the leg and forearm. 
Dorsally the supply of the tibial digits, wholly or partly, by the 
anterior tibial derivation from the posterior tibial nerve in this 
and other animals, would scarcely have been expected from anything 
in the construction of the hind limb. The anterior tibial nerve is in 
the fore limb represented in Cryptobranch by the branch of the 
median passing dorsally to join the radial and inferior musculo-spiraL 
The fibular digits are supplied by the musculo-spiral (peroneal) 
of the sciatic and the ulnar digits by the inferior musculo-spiral 
branch of the median: the only difference being that the nerves 
to the outer digits in the fore limb keep company with the main 
nerves of the limb (median or median and ulnar) longer than their 
correspondents in the hind limb keep company with the sdatia Or, 



THE CBTFTOBRANCHUS JAPONICUS. 55 

• 

it might be said, the mubculq-spibal in the hind limb detaching 
iteelf from the main trunks carries with it the dorsal nerves of 
the fibular digits. In ourselves the dorsal nerve of the ulnar digits 
— the 'iKFERiOB musculo-spibal' as it may be called — ^keeps company 
with the ULNAR nesve to the middle of the forearm, and the dorsal 
nerves of the fibular digits (the external saphenous) is derived partly 
from the musculo-bpibal or peboneal, and parti/ from the pop- 

UTBAL. 

The chief differences from the corresponding nerves in Man, in 
addition to those juut mentioned, are (1) the deeper position of the 
digital branches in the palm and sole beneath, instead of superficial 
to, the flexor tendona In the fore limb the branches to the ulnar 
digits are derived from the deep palmar branch of the ulkab instead 
of from a superficial branch ; and those of the radial digits come frt>m 
the IHTEBOSSEOUS which (in Cryptobranch) forms the main trunk of 
the MEDIAN, instead of coming off near the elbow by a branch which 
separates frt>m the interosseous upon the flexor pbofundus, and the 
8i2e of which causes it to be regarded as the main trunk. (2) In the 
hind limb the extebnal popliteal is, in Cryptobranch, like Uie ulnab, 
continued to the digits; whereas in Man the course of the nerves to 
the fibular digits is not along the fibula, with the nerve which accom- 
panics the peroneal artery, and which is the representative of the 
ULNAB nerve, but with the POSrrERiOR TiBiAii, which is the representa- 
tive of the MEDIAN, till they reach the foot, where they separate, and 
form the external plantar nerve. 



An interesting feature in the nerves of this animal and its allies 
as compared with those of higher animals is the simple structure of 
the BBACHIAL and cbubal plexuses, which may be associated with 
the imperfect s^^entation and specialization of the action of the 
limb-muscles. The office of these plexuses, we may conclude, is to 
ensure that harmony and those nice balancing modifications in the 
degi-ee of action of the muscles — the opponents as well the co-opera- 
tives — upon which the adjustment and proper direction of the move- 
ments of the limbs depend. This, it may be inferred, is effected by 
such an arrangement of bundles in the plexuses that a certain number 
of the filaments of any nerve presiding over one muscle, or set of 
muscles, are distributed among the nerves destined to each of the 
other muscles or sets of muscles ; so that an impression transmitted to 
one muscle or set of muscles is in duly ordered de^pree communicated 
to each of the other muscles. Where, consequently, the muscles are 
numerous, the interchanging branches between the nerves requisite 
for this purpose will be numerous and form a complicated plexus; 
and where the muscles are few the plexus will be proportionately 
simple. The same object may be attained by intercommunicating 
filaments between the nerves in the distal parts of the limb, or by two 
or more passing to the same muscle. Thus the ninth and the 
cervical nerves intercommunicate before supplying the omo-hyoid in 
ourselves, while the stemo-mastoid and Pmpezius are each supplied by 



56 PROFESSOR HUMPHRY. 

branches of the spinal accessory and of the cervical nerves passing 
separately into them; and the cominnnicating branches above described 
between the obturator and sciatic, and between the median, radial, 
and posterior ulnar at the back of the forearm, indicate that the 
offices performed by the proximal plexuses in ourselves are, to a 
greater relative extent, performed by distal plexuses or oommuni- 
cations in the CryptobranclL 

The course which the nerve-current follows, like that of the blood- 
current, is, in each case, probably regulated or greatly modified by 
convenience; and a nerve in its way to a particular muscle takes 
usually the most direct course, and may attach itself to one set of 
nerves or another, according as either lies more across its path. I 
have already alluded to the difference in rotation of the fore and hind 
limbs as a cause of difference of distribution of the nerves in the two. 
The rotation in each limb attains its maximum in ourselves, and 
causes the divergence of the nerves in the two limbs to be greater 
than in the Cryptobranch. For examples; the extensors of the 1^ 
and of the forearm are both in that animal chiefly supplied by nerves' 
passing behind the respective girdles ; in Man the nerves to the ex- 
tensors of the forearm pass behind the scapula, whep^ those to the 
extensors of the leg pass in front of the pelvis. — rin Cryptobranch, 
the nerves to the muscles on the dorsum of the scapula and to the 
pectoral pass behind the shoulder, and those to the muscles on the 
dorsum to the ilium and to the gracilis pass behind the hip ; in Man 
the scapula being thrown as well turned more backwards, the nerves 
(pectoral and suprascapular) to the corresponding muscles in the fore 
limb pass in front of the shoulder, while those in the hind limb 
(obturator and gluteal) pass behind the hip. — In Cryptobranch the 
nerves on the palmar and plantar aspects of the two limbs below the 
elbow and knee present scarcely any difference. In Man the nerves 
to all the digits of the hind limb proceed together in the posterior 
tibial nerve as far as the ankle ; the companion of the peroneal artery, 
which is the representative of the ulnar nerve, is a mere muscular 
branch; whereas, in the fore limb, the nerves to the ulnar digits take 
their course down the forearm in the ulnar nerve quite separate from 
the nerves to the other digits, which are boimd together in what we 
call the median nerve. These points indicate, as do the variations 
observed in this and other animals and in man, that nerve-course is 
somewhat too arbitrary or too much regulated by convenience in each 
instance for us to be able to rely upon the disposition of nerves as 
at all sure guides to the discernment, in difficult cases, of the homo- 
logical relations of muscles and other structures. 



THE CRYPTOBKANCHUS JAPONICUS. 57 



DESCRIPTION OP FIGURES. 

Thej are all representations of dissections of tke dyptobrancli 

unless it is otherwise specified. 

Fig. 1. Transverse section through tail. F.^ fascial investment 
covering the part, ensheathing the ventral and dorsal lateral muscles 
and sending down lamime to the vertebral processes. D.f.^ dorsal 
&trma«8 in triangular space between fieiscial laminee on either side 
descending to dorsal spiDOS and superficial lamina passing across. 
v./., and Z./, veDtral and lateral fat-masses similarly situated. S, tf,y 
supra-lateral fat-mass with posterior spinal nerve curling from beneath 
it. The infra-lateral fat-mass is similarly situated. 

Fig. 2. Muscles of the left side of the tail, hinder part of 
abdomen, and dorsal aspect of left thigh. 2>., dorsal muscle of tail. 
F., ventral muscle of tail. The lateral septum is seen between these 
two. /. c., The iliac part of ventral muscle (Ilio-caudal) passing over 
the ilium (/.) and continued into the internal oblique (/. 0.). E, 0., 
external oblique which has been partly divided and refiected up- 
wards and downwards to expose the internal oblique and trans versa- 
lia. E, (y.y the part of external oblique attached to the spine of the 
pubea and extending upon the hip between {Or.) gracilis and 
{Ped,) pectineus. ^V., tiunsversalls exposed by reflection of ex- 
ternal oblique and removal of part of internal oblique. Py., pyramid- 
alis. //., iliacus. GL n, gluteo-rectus. B, biceps. R, t., rectus 
intemus. T, a,, tibialis anticus. Ext. d., extensor digitorum. P., 
peroneus. 

Fig. 3. Right side of hinder part of abdomen, of tail and right 
bind limb shewing the caudal muscles and the extension of the ex- 
ternal oblique into the fascial tissue on the dorsum of the thigh. 
D.J dorsal muscle of tail. 7. c, ilio-caudal. C,/,, caudo-femoxal. 
C. cr,^ caudo-crural. Isch, c, ischio-caudal. E. 0,<, external oblique 
reflected from around the ilium to show its connection with the ilium 
(p. 13). E, O'^.j external oblique of the left side inserted into the 
edge and horn of the prepubic cartilage. 

Yi^f, 4. Interior view of the muscles on the left side of the 
under surface of the back of the abdomen with the nerves of the 
brachial plexus. F., F., right sides of the bodies of the vertebrce 
uncovered. /. v, inter-vertebral substances. S, v. jB., sub- vertebral 
rectus. D. C, depressores ooetarum. JV., transversalis. /. 0,, in- 
ternal oblique seen in shadow through the hole cut in the transversa- 
lis. ilT., Nerve lying between transversalis and internal oblique. 
In the upper part of the drawing are two separate portions of the 
internal oblique which arise from the fourth and fifth ribs. (7., 
coracoid. P. C, pre-ooraooid. S,, serratus. Tr,, trapezius. L. S,, 
levator scapnlie. C,/., constrictor fitudum. F,, Vagus nerve. 



58 PROFESSOR HUMPHRY. 

O.h.N., omo-hyoid nerre coming from the third cervical nerve. 
6, the sixth cervical nerve, the hindmout contributor to brachial 
plexus, indicated by a dotted line where it is covered hy the trans- 
yersalis. It is seen in front of the transversalis joining the .fifth 
cervical nerve. 

Fig. 5. Under surface of fore-part of trunk, neck and shoulders. 
E,0.^ external oblique passing into (P.) pectoraL The right pectoral 
and external oblique have been reflected with the coracoids showing 
/. 0., internal oblique attached to upper plate of (iS^.) sternum (p. 15); 
the lower plate is exposed beneath it. Scy scapula. C7., coraooid, 
P, c, pre-coracoid. C, b. br., coraco-brachialis brevis. C. o., coraoo- 
olecranalis. 0. h,, omo-hyoid. F, c. 5., precoraco-brachial. £p, c. 6., 
epicot'aco-brachial. />. «., dorsalis scapulse. B. jET., Basi-hyoid with 
cartilaginous nucleus, behind its middle, in the branchio-hyoid liga- 
ment. C. H,y comua of the hjoid. E, C, entoglossal cartilage. 
£^yB\ first branchial arches with the copula ((7.) between them. 
B*, second branchial arch. S.cG.^ subcutaneus colli divided % and 
reflected to either side and seen passing uader and attached to the 
ramus of the jaw. M, ff,, the mylo-hyoid. D. M., depressor mandibulse. 
D. digastric. S. If,, foremost part of internal oblique or stemo-hyoid 
passing to ((7.) copula and entoglossal cart. O, 6., genio-branchial. 
B. J7., branchio-hyoid. /. M, iuter-mandibular. 

Fig. 6. Deep view of under^surface of neck with right fore-limb. 
S. V, R,f subvertebral rectus passing to base of skull. L. «a, levator 
scapulse. Tn, trapezius inserted between pre-coracoid and scapula. 
J).8C., dorsaUs scapuke. S, C, subcutaneus colli. Cr./.^ constrictor 
faucium. D, Jf., depressor mandibulse. Z>., digastric. C, coraooid. 
P. 0,9 pre-coracoid. ' 

Fig. 7. Diagrammatic representation of abdominal muscles more 
particularly of the external {K 0,) and internal (/. 0.) obliqae and 
the rectus. E. 0. E,, the external layer of rectus formed by the 
fibres of the external oblique. /. 0. R.y the deeper layer of rectus 
formed by the fibres of the internal oblique. TV., transversalis. 

Fig. 8. Muscles t>f tail and left hind-limb : the latter has been 
turned up so as to shew the plantar surface. Z>., Dorsal, and T., 
ventral muscles of tail separated by the lateral septunL /. c, ilio- 
caudal. Isch. a, Ischio-caudal. C.f»<, caudo-femoreJ. C. cr., caudo- 
crural. C.p.y caudo-pedal. /.y!, ilio-femoral, or bicepa -^.^i 
femoro-fibular, or short portion of biceps. P., pubea. Gr^ giucilia. 
iSif., semitendiuosus. T.a., tibialis anticus. FL cL 8., flexor digitonun 
sublimis. 

Fig. 9. Plantar aspect of left hind leg and foot. C. p^ caudo- 
pedal. i7. ^., flexor tibisB (aemitendinosus, &a). FLp.^ flexor maas 
of foot. FL d. 8,y flexor digitorum sublimis divided and reflected 
upwards and downwarda Fr,p., pronator pedis, or part of flexor 
mass inclined to tibial side of foot with its superficial fibres running 
into under-surface of flexor dig. subl. Fl. cL p., flexor digitorum pro- 
fundus. N,j nerve to sole lying on FL dig. Abd,y abductor minimi digitL 



THE CRTPTOBRANCHUS JAPONICUS. 59 

Fig. 10. Miudea and nerves on plantar aspect of left hind limb, 
/.y internal surfaoe of ilium with fibres of internal oblique passing 
to and j&om it. Cf,^ caudo-femoraL (7. CT.y caudo-crural. C. p,j 
cando-pedal joined hj portion from ischium. Fl.e,^ flexor cruris. 
Gl, r.y gluteo-rectus. B, biceps. F. f.^ femoro-fibular. S.^ sciatic 
nerve passing beneath ilium and sending nerves to above muscles. 
P.y peroneal nerva E. P., exteraal popliteal /. P., internal pop- 
liteal. Pr.^., pronator pedis. /7. ^.je>., flexor digitorum profundus 
Fl. d. 8,y flexor digitorum sublimis, of which the lower part has 
been removed. 

Fig. 11. Muscles and nerves on plantar aspect of leg and foot. 
Fl, c, flexor cruris. a,a.a,j flexor sublimis, divided and reflected 
upwards and downwards (p. 26). FLp., flexor profundus. Pr.p.f 
pronator pedis. £. F. iiT., external popliteal nerve. /. P. ilT., in- 
ternal popliteal nerve. 

Fig. 12. Deep muscles of sole. FLd. 8., fiexor digitorum sub- 
limis turned up^ and the lower part removed, shewing the cut edge of 
fibres which passed into Fl. d. s. Fl. d. p.j flexor digitorum pro- 
fundus divided, the lower part removed. Fr.p., pronator pedis 
rising at the lower part to join Fl. d. s. T. m.y tarso-metatarsals. 
M.p.y metatarso-phalangeL JPA. phalangeL /., interosseus. Abd.^ 
abductor minimi digitl 

Fig. 13. Back of leg and sole of Menobranch dissected to shew 
the flexor digitorum profundus nearly divided in two, the tarsal car- 
tilage intervening between its crural and its pedal parts. FL d. «., 
flexor digitorum sublimis. Fr. p., pronator pedis. 

Fig. 14. Deep dissection of plantar aspect of right hind leg 
shewing Pn ^., pronator tibin. FLcy flexor cruris reflected. Cp., 
caudo-pedal FLd.8.y flexor sublimis reflected upwards. Fr.p., pro- 
nator pedis reflected; below it is blended with flexor subl. FL d.pr.^ 
flexor profundus. T., lower end of tibia. The fibular and posterior 
tibial nerves are shewn, the latter passing in front of pronator 
tibiae. 

Fig. 15. Muscles on dorsal aspect of left leg and foot. C, con- 
dyles of femur. /. if., internal rectus. T. a., tibialis anticus passing 
to tarsus; T.c^. tibialis anticus inserted into tibia. GLr.^ tendon 
of glnteo-rectns passing to lower end of tibia between extensor 
digitorum and tibialis anticus ; E. d., upper part of extensor digi- 
torum turned to one side. E.c^., portion of extensor digitorum 
joining peroneus (P.)' E. d'\y extensor digitorum reflected down- 
wards. S. p., supinator pedis. E. d. br., extensor digitorum brevis, 
E. d. br^.y slip from extensor brevis to supinator pedis. Abd, ab- 
dnctor minimi digitL 

Fig. 16. Yiew and letters similar to preceding. /., interossei 

Fig. 17. Muscles of left forelimb, with the radial and inferior 
muscnlo-spiral nerves seen in the back of the forearm. C, coracoid. 
P. (7., pre-coracoid. S.y scapula. E.p. c, epicoracoid muscle. P. c. ft., 



60 PROFESSOR HUMPHRY. 

pre-coraoo-brachialis. B. biceps, detached from the scapula and turned 
back to shew (C. b, br.) the coraco-brachialis brevis. C, b. Z., coraco- 
brachialis longus. Pect, pectoral. 2V., trapezius. L. s., levator scsr 
pulie. D. 8.f dorsaJis scapulae. Br, a., brachialis anticus. JExt. c. r., 
extensor carpi radialis. Uxt. d. «., upper and lower parts of Ext. d. 
Bublimis, of which the greater portion has been removed. £xi. e. u.j 
extensor carpi ulnaris. Sup. Ly supinator longus. Sup, br,, supinator 
brevis. Sup, m., supinator manius. East d, br., extensor digitorum 
brevis. 

Fig. 18. Muscles of right forelimb. C7., ooracoid with {Ep, c, b,) 
epicoraco-brachialis. B., biceps. H,, humerus. C. b, I,, coraco- 
brachiaJis longus, with divisions to biceps, to ulna, and to condyle 
of humerus. Pect,, pectoral divided and thrown outwarda Br, a., 
brachialis anticus. Fl. c, r., flexor carpi radialis. Pr, L, pronator 
teres perforated by the nerve to (Fl, d, a,) flexor digitorum sublimis, 
which has been divided and reflected upwards and downwards. 
^/. c. ft., flexor carpi ulnaris. Fl,d,p., flexor digitorum profundus. 
Pr, q.f pronator quadratus. Pr. m., pronator man^B. B, radius. 

Fig. 19. Muscles and nerves on palmar aspect of right forearm 
and hand. Fl. d, a., flexor sublimis digitorum divided and reflected 
upwards to the humerus and towards the ulna and downwards at its 
attachment to the digits. Fl,d,p,, flexor digitorum profundus ra- 
diating to the digits with the carpal cartilages appearing through it. 
Pr, m,y pronator maniis divided and reflected. Pr, t,, pronator terea 
Fl. c, r,, flexor carpi radialis. Br, q,, pronator quadratus. P pha- 
langeus. U,, ulnar nerve and branches. Jf., median nerve. 

Fiir. 20. Muscles and nerves on the upper part of the head. 
T., temporal muscle of right side; on the left ?de the npper part of 
the muscle has been removed leaving only the part near the inser- 
tion, if., masseter. On the left side the upper part has been re- 
moved (exposing the suspensorium with the dentary nerve in front 
and the seventh nerve behind); and the lower part has been turned 
back to shew its insertion into the lower jaw apd the dentary 
nerves passing into the jaw. On the right side a branch of the thii'd 
division of the AM is seen emerging from beneath the master and 
joining the seventh. 7a. Conjoined branch of 7th and 5th — the sub- 
mental nerve. In the left orbit are seen the ophthalmic nerve 
near the inner part of the orbit giving off the ciliary nerve and the 
supramaxillary, with the optic crossing fix>m the inner side of the 
orbit over the latter. 2f., hinder end of the hyoid cornu with {St) 
styloid cartilage between it and the jaw. Z>., dorsal muscle ex- 
tending to head. 

Fig. 21 represents a piece of the side from which the lateral 
musole (L,) has been partly removed to shew one of the spinal nerves 
emerging from the vertebral foramen just behind the rib (E.) and 
dividing into anterior or intercostal or ventral (F.) and posterior 
branches. C, cutaneous nerve from the latter. The posterior 
branches pass in front and behind the articulating processes to the 



THE CRTPTOBRANCHTJS JAFONICUa 61 

dorsal part of the lateral muscle. The anterior branch nms along 
the hinder border of the rib. I^ inscription in the donal muscle. 

Fig. 22. View of the brain and cerebral nenres in situ, the upper 
part of the skull and the meninges with the fine areolar tissue which 
occupied the large interval between the dura mater and the skull 
having been removed. 1. The olfskctory nerve expanding into the 
c^iebral hemisphere (Schmidt^ G. and Y. der H. judged that there 
was A cavity in the henuspbere. Tn mj specimen the part was too 
soft for this to be decided). The hjpophysb was very large and 
occupied a distinct digital fossa on the upper surface of the basis 
cranii. There was also a Pineal Gland but much smaller. 2. The 
optic nerve passing from beneath the hemisphere. The two optic 
nerves are quite separate from one another, and do not cross, each 
running direct from the hemisphere to the orbit of the same side. 
3. The third nerve {motor oeult) passing from beneath the hemi- 
sphere. 4. The fourth nerve passing from the strand at the side of 
the third ventricle. The third and fourth nerves escape through 
small foramina in the dura mater and apparently accompany the 
ophthalmic part of the fifth. 5. The fifth nerve passing from the 
side of the forepart of the brain bounding the fourth ventricle : it 
expands into the Gasseiian ganglion as it enters the dura mater. 
7. The seventh nerve passes from the side of the back of the brain 
bounding the fourth ventricle and divides into the portio dura and 
the auditory part. 8. Two nerves passing from the medulla oblongata, 
meeting as they enter a common hole in the dura mater and skull, 
and forming the eighth nerve. The anterior of the two is probably 
the glosso-pharyngeal, and the small nerve seen joining the vagus 
from behind Lb probably the representative of the spinal-accessory. 
They are so regained by Schmidt, G. and Y. d. H., who also describe 
a nerve which they traced through an anterior condyloid foramen as 
the 9th or hypoglossal nerve. We could not discover any represen- 
tative of the 9th nerve in the cranial cavity. 



ON THE COMPOSITION OF THE CARPUS OF THE 
DOG. By Pn)/e«ar W. HL Floweb, F.RS. (Read at the 
Meeting of the British AsBociation, at Edinburgh, Aug. 7, 

1871.) 

The number of the elements fonning any part of the skeleton 
in the larger number of animals and therefore regarded as 
normal or 'typical', may be exceptionally reduced in certain 
species, either by absolute suppression or by the union of two 
or more element& 

This union may be brought about either by (1) coalescence, 
where a single part is formed by the fusion of two or more 
elements developed separately in the embryo; or (2) conna- 
Bcence, where a part which (as £Gur as can be traced) has been 
single from the first, in one individual, represents by its form, 
situation and connection with surrounding parts, two or more 
distinct elements in another individual 

An example of coalescence in the carpus, is seen in the genus 
Salamandra, in which, according to Oegenbaur's observations\ 
the intermedium and ulnare are distinct in the new-bom, but 
united in the adult animal. 

On the other hand, the undforme of the mammal's carpus 
which represents the two carpal bones of the ulnar side of the 
distal row of reptiles, is, as far as is at present known, an 
example of connascence; for in no mammal has this bone been 
found to have origin in two distinct centres of ossification, or in 
separate cartilages. 

This distinction is important, although it is obvious that 
it can only be employed in cases in which our knowledge of the 
development of the part is complete, and a gradual conversion 
of the less into the more perfect condition of union in successive 
generations and by various intermediate or transitional condi- 
tions is perfectly conceivable. 

In the dog and other animals of the order Camivora, as 
is well known, a single bone represents, by its position and rela- 

^ Untertuchungen Bur vergleichenden AnatomU der Wirbelthiere, Itot Heft. 
Carpui vnd Tamu, 1864, p. 5, and PL I. figs. 1 and 8. 



.COMPOSITION OF THE CABPUS OF THE DOG. 63 

tion to suntmoding stnicturaa, the radiate and interme(Uum of 
the typical carpus, more commoDly called in England the 
scaphoid and the lonar. In this, as in other similar cases 
unoDg the Rodentia, Insectivora, &c. researches in the embryo- 
nic condition have not hitherto, according to Gegenbanr', shown 
any true coalescence of distinct elements. 

The Camivora, moreover, have always been included among 
the orders in which the ot centrale (a bone found in monkeys, 
many Rodents, Insectivores, &c.) is wanting as a distinct ele- 
ment*. This bone, previously described as a dismemberment 
of one of the more constant bones of the carpus, is considered 
by Gegenbaur as a true primitive element of that region. He 
adds : " But what takes place in those cases in which it is 
wanting, has not yet been discovered. My endeavours in the 
examination of embryos, both of man and of many nuunmals, to 
obtain any indication have all been in vain, and I can only say 
that there are no grounds for the acceptance of its connection 
either with the nuifrnum (Cuvier) or with the scaphoid (Owen)*." 

A recent examination of the natural skeleton of a dog, 
about six weeks old, which I had prepared for the museum of 
the Collie of Surgeons of England, has fortunately thrown 
light upon both these questions; for I find that at this age, the 



Dnsal fOTtioe ot right e»rpai ol 7oiiiig Dog, Ho. 4419 d DiteoL B«riM, Mac 
Boy. Coll. Suig. Ixmdrai z. S. e mmeiform or ulnare, I loaax or iniemediuM, 
te mphoid or ratliaU, ee oentral, tm trftpezinin, (li tnipeEOid, m magnnm, u nn- 
eifonn, p pisiform, I-^T metMupali. 

I Op. tit. p. 44. 

■ QagraUar, op. ett. p. 50. W. H. Floww, Iturodueliom to the OiUologj/ of 
tkt MamHaHa, p. StiS. 
* Op. eit. p. 60. 



64 PROF. FLOWER. COMPOSITION OF THE CARPUS OP THE DOG. 

80-caIled scapho-lunar bone, though well ossified, consists not 
only of a perfectly distinct scaphoid and lunar, but also of a 
third piece evidently corresponding to the os centrals of the 
typical carpus. 

The two former are divided by a vertical suture in the 
usual situation, the scaphoid being rather the larger of the two. 
The cerUrale is a small wedge-shaped bone, with its broad end 
towards the dorsal surface of the carpus, where it appears in 
the form of a transversely elongated lozenge, articulating above 
(or proximally) with both scaphoid and lunar, and below (or 
distally) with the trapezoid and magnum, fitting into the groove 
between the two. It thus lies altogether between the first and 
second row of carpal bones. As it recedes from the dorsal 
surface it narrows to a point which does not extend so far as to 
appear on the palmar aspect of the carpus. It is rather smaller 
and more flattened than the corresponding bone (which remains 
permanently distinct) in a monkey, but has precisely similar 
connections. The disposition of these structures is exactly the 
same on both sides of the body. 

Although it is desirable that this observation should be 
repeated in other Camivora, it appears to me important as 
proving that in the dog at least, neither the radiale (scaphoid),' 
interTTMdivm (lunar), nor the centrals are suppressed, neither 
are they connate, but they are all developed independently 
and afterwards coalesce to form the so-called ' scapho-lunar ' 
bone, and further, because it lends additional weight to the 
view of the nature of the os centraie advocated by the distin- 
guished anatomist of Jena. 



DESCBIPTION OF THE BRAIN OF AN IDIOT. By 
S. Messenger Bradlet, F.RC.S., Lecturer on Hwman 
and Comparative Anatomy, Royal School of Medicine and 
Surgery, Manchester, Plate Y* 

General History. The idiot from whom the brain was 
taken which is here described, died from bronchitis in the 
Withington Workhouse, Manchester, of which he had been an 
inmate for fifteen. years, at the age of 35. His parents, who 
are people of average intelligence, on being interrogated, 8tate4 
that there was no other instance of idiocy, or yet of epilepsy 
or insanity in the family. It was difficult to correctly ascertain 
what height he was, as he always lay, or sat, coiled up into 
a ball, with his arms and legs acutely flexed upon themselves ; 
he certainly would not have exceeded four feet, however, if it 
were possible to have stretched him out. 

At death he weighed only 60 pounds; he was always 
extremely feeble, being indeed little more than skin and bone. 
His attitude in life was remarkable ; he sat throughout the day 
huddled together like a tailor, upon a little bench, continually 
swaying to and fro, and constantly uttering a low moaning 
kind of note. 

Senses and Intelligence, Both taste and hearing were pro- 
perly developed, but he was blind from his birth. For some 
years preceding death there was opacity of both comese, with 
atrophy of the eyeballs. Up to the age of nineteen he never 
spoke, but he subsequently learnt to say a few words, which he 
would repeat parrot-like, when the cue was given him. They 
consisted in the repetition of much such a formula as the 
following : " Jaunty (his name was Jonathan Gorman, but he 
was always called Jaunty), good boy, Christians awake, happy 
mom, four and twenty blackbirds baked in a pie. Mother'^ 
(pronounced Mudd-der). This with the addition of some pro- 
fane expressions completed his stock of learning. He scarcely 
ever varied his formula, although occasionally, like " Grip," he 

VOL, VI, 5 



66 MB BBADLET. 

lost his way and mixed up his oaths and nursery rhymes in 
singular confusion. 

His habits were filthy, and his passions violent : he never 
recognized any one, and always bit and tore at the man who 
daily carried him from his seat on the form, or chair, to the 
watercloset^ where he made it his invariable rule to eat his 
taeals : this strange custom he persisted in for many years 
before his death. He was not paralysed, but was quite in- 
capable of any attempt at locomotion. He was equally unable 
to dress himself, or indeed to perform any of the functions 
of life unaided, beyond carrying his food from his plate to hid 
mouth with his fingers. 

He was very restless when lying down, and in tossing 
about from side to side he sometimes jerked his arms or legs 
against the edge of the bed, and when he did so, he invariably 
fractured a bone. These fractures always united, but owing to 
his extreme restlessness, it was found impossible to keep the 
firagments in apposition by any arrangement of splints or ban- 
dages, and in consequence of this almost incessant movement^ 
the bones united at the most extraordinary angles ; e.g. the 
femora and humeri, which were all fractured at one time or 
another, were so acutely bent upon themselves during the pro- 
cess of union, that the upper and lower fragments assumed an 
almost parallel position to each other. After death nine im- 
portant fractures were counted, and several of less moment. 
In addition to these it was found that seven of the ribs had 
been broken at periods more or less remote. 

Neither the expression of the face, when asleep, nor the 
external shape of the head, conveyed the notion that the man 
was idiotic. The face, which was quite the best part of him, 
was rather intelligent-looking than otherwise*. 

1 The flkoll was bmchyoephalio, tMafiuing 6*5 aerois tiie vertex from rid», 
and 6 inohefl from before baokwards. I do not however think that this fact off 
braohycephalism is in itself of modi yalne, or that the olassifioation whi<^ 
Betzias fotinded npon Uie rehitive lengih and breadth of akuUs is a good one. 
I am at present engaged with Dr Lyson in investigating the varieties of English 
Omnia, and, withont anticipating the paper we hope to publish on this aobjeoty 
I may here state that we meet with extreme cases of doheoeephalism and 
braohycephalism amongst the English residents of our own town. Huxley in his 
remarks upon the ' Neanderthal ' skull oomes to much the same eondnsim 
whidi we have arrived at, when he affirms that " oranial meaearements alone 
afford no safe indication of race". 



DESCBIPnOH OF THfl BRAIH OF AN IDIOT* 67 

Weight and general appearance of the hraif^ and of its eomr 
ponent parts. After removal of the membranes the brain 
weighed 28 ounces^ which was made up in the fbllowing 
manner': 

The cerebrum 22*8 oz« 

TU^ «o«.^^o^l„«, ffiigtt hemisphere 25 ... 
TLne cerepellum ix ^l i. • ^ e%u\ 

iLett hemisphere 2*0 ..^ 

The pons and medulla '7 ... 

280 oz. 

The most striking peculiarity here is the disproportion in 
weight between the two cerebellar hemispheres, but in spite of 
the deficiency in the left half, it is noteworthy that the propor- 
tion to the cerebrum of the entire cerebellum, 1 to 5*5, is, as is 
usual in the brains of idiots, considerably higher than it is in 
healthy brains, where it is 1 to 7. When first removed from the 
calvarium the cerebellum was entirely concealed by the cere- 
brum, and at first sight the general appearance of the encephalon 
conveyed the idea of a small but normally developed brain. 

On section the grey matter was found to average *14 inch 
in thickness, and appeared paler than usual, but microscopically 
the cells presented a normal appearance. The sulci averaged 
half an inch in depth, but were deeper at the front than at 
the back of the bzain, being '6 inch in the frontal, and only 
'4 inch in the occipital lobes. The pons varolii measured 
"S i^ch in thickness. 

The cerebellum afforded the following measurements :— * 

LeA hemi^here of cerebellum : 

From before backwards 2*25 inches. 

From side to side '9 

From above downwards 1*5 

l^ht hemisphere of cerebellum : 

From before backwards 2*5 

From side to side 1'2 

From above downwards 1*5 



... 



... 



a * . 



... 



. • • 



1 nil is not at aU an aitremelsr light 1»aixi for an idiot, and Msarody biringi 
it into the eategoiy of micio-eephalic brains ; inetanoeB being recorded of brains 
ireig^xing reapectirely 16-6 oz. (Wagner's Vorstudien), 18 125 oz. {Traiu, ZooL 
Soc. I. 343), 10 oz., and 510 09. CTr<»n#. PAiJ. Soc. 1864, Fart IH. p. 537). 

6—2 



68 MR BRADLEY. 

Fissures, Lobes and donvohftions seen on the superior and 
outer a^ect of the brain. 

Fissure of Sylvius (8) — Right side. The bonzontal limb 
was observed to extend -within a very short distance of the 
intra-parietal sulcus {IP). The ascending limb (S^ ran into 
the second' frontal convolution (2 — 2). Left-side. The hori- 
SEontal limb did not run so far on this side as on the other by 
an inch. 

Fissure of Salando (B). This fissure was situated almost 
midway between the frontal and the occipital lobes, and con* 
sequently decidedly more anteriorly than is the case in well- 
developed brains* This is explained by the imperfect, post- 
natal, development of the frontal lobes. Right-side. Did not 
quite reach the horizontal limb of the fissure of Sylvius (8)^ 
so that the ascending frontal (4 — 4) and parietal (5 — 5) con- 
volutions were continuous. Left-side. A similar dispositloa 
was met with on this side, but it extended quite into the longi* 
tudinal fissure. « 

External Parietooccipital Fissure (PO). Right. Swept 
abruptly backwards, and then ran forwards and outwards for 
three inches towards the Parallel Sulcus (P) into which it 
merged. In consequence of this arrangement the first annec- 
tent convolution (a — a) was apparently not present, and the 
second 03 — )3) was removed to a greater distance than usual 
from the longitudinal fissure. ' Left. Of the usual extent, it 
cut the upper surface of the brain for the space of about half 
an inch. 

Intra Parietal Fissure (IP)^ Right. The ascending Kmb 
ran parallel to the fissure of Rolando {Rj as far as the longitu- 
dinal fissure: the horizontal limb (on the contrary) was ex- 
tremely short. The ascending limb was separated from the 
fissure of Sylvius by the supra-marginal lobule (A). LefL 
Ascending limb not so extensive as on the right side, but hori- 
zontal limb somewhat longer. 

Parailel Fissure (P). Right. Ran backwards into the exter- 
nal parieto-occipital fissure (PO) as before mentioned. The 
parallel fiissure and the fissure of Sylvius were, however, not 
joined. Left Parallel fissure separated firom external parieto- 
occixHtal fissure by a distance of 1'5 inches. 



J 



DESCRIPTION Of THE BRAIN OF AN IDIOT. 69 

The Lobes and Convolntions. The frontal convolutions of 
both hemispheres were non-symmetrical and fairly developed, 
though relatively small; they all sprang from the ascending 
frontal convolutions. Supero Frontal (8F) and Infero Frontal 
{IF) sulci on both sides interrupted by secondary convolutions. - 
The third left frontal convolution was very well developed. 
Orbital convolutions presented nothing remarkable. The ascend- 
ing Frontal (4 — 4) and ascending Parietal (5 — 5) convolutions 
were large and partially cut by secondary sulci. On the right 
side the lobule of the ascending parietal convolution (5' — 5') 
was connected with the angular gyrus (6 — 6), and ran back to 
the external parieto-occipital fissure {PO), which it crossed by a 
small bridge of white matter (the position of which is indicated 
by the mark *) lying '2 inch below the surface, and hence con- 
cealed from sight until the sulcus was stretched apart. In this 
manner the parietal and occipital lobes were united by a first 
annectent convolution (a — a) which was hidden from superficial 
observation. On the left side the lobule of the ascending 
parietal convolution (5' — 5') was separated from the angular 
gyrus (6 — 6) by a shallow sulcus, but was connected in the 
ordinary manner with the occipital lobe by a superficial an^ 
nectent convolution {a — a). 

Annectent Convolutions {a — a), (fi — ^/8), (7 — 7), (S — S). 
Bight-side. The first was deeply placed, as before stated, and 
was small. The second and third were also small, and almost 
separated from the parietal lobe by the near junction of a deep 
transverse fissure with the parallel fissure (P). Though small, 
they were present however. Left-side, Well and normally 
formed. 

Lobtde of the Supra-Marginal Convolution (A). Bight side. 
Small, joined angular (6 — 6) and ascending parietal convolution 
(5 — 5). Left side. Kather large and more quadrilateral : united 
the same convolutions on this side as on the other. It is im- 
portant to note the presence of this convolution in both hemi- 
i^heres. In Professor Marshall's cases of the micro-cephalic 
brains of two idiots (reported in Phil. Trans, for 1864, Part III.) 
it was nowhere present. Both by him and by Qratiolet it is 
i^garded as, perhaps, the most characteristically human of all 
the convolutions. 



70 KR BBlDLEt« 

Angular of Bent Ccmvolution (6-^6). Bight. Compl€% di- 
vided into numerous secondary convolutions, and continuous 
with the second annectent convolution 03 — P). Left, Markedly 
simpler, but also continuous with second annectent convolution 

08— i8). 

Fieeures, Lobes, and Ckmvolutions of Internal a/nd Inferior 
Surfaces of Ger^yrum. 

The OaUoso-marginal, the Oalcarine, and the Pentate fissurefi 
preseuted no peculiarity worthy of note. - 

CoUaterai Fissure. Bight. This important fissure could 
not be said to exist at all on this side; but instead of this it was 
found that a branch of the parieto-occipital fissure joined a deep 
sulcus, one inch in depth, which ran at right angles to it, trans* 
Versely across the brain, apparently cutting off the parietal 
from the occipital lobes. This transverse fissure (which was 
notably the deepest fissure in the entire brain) ran within *2 inch 
of the parallel sulcus (P); if this junction had actually taken 
place, the parietal and occipital lobes would have been com- 
pletely disjoined; but, as has been described, the annectent con- 
volutions though small were present^ and crept into the narrow 
space between the two sulci*. 

The GaMosai and marginal convolutions were well developed : 
the occipital lobule was smaller than usual on both sides, but 
not otherwise remarkable. 

Isla/nd of Beil. Quite concealed on both sides. Symmetri- 
cal in the two hemispheres, and consisted of three perfectly 
smooth, small, radiating convolutions. 

The Temporo-sphenoidai Lobe was well developed, &nd 
nearly symmetrical on the two sides. On the right side the 
external temporal convolution was very uhUdUAlly thin. 

The Corpora StrioM were rather smaller than the Optic 
Thalamii The ventricles measured 1*25 inches from side to 
side. 

The Corpora Quadrigemina, although composed of their 
usual elements, were extremely small, being not more than half 
the usual size. 



1 Gratiolet has fignred a somewhat similar saletiB as existing in the bnin of 
a porpoise ^Lenret et Gratiolet, Atlas de Vanat. eomp. 1857. Plate XI.), and in 
the hrain of a six months* foetos (loe. eit. Plate XXX. Fig. 1.). 



DESCRIPTION OF THE BBAV OF AN IDIOT. 71 

Certidlvm. The yermiform process was imperfect: the 
fiodvle and uvula were present, but small; the pyramid and 
commissura brenis were entirely undeveloped* The weight and 
measurements of the cerebellar hemispheres have been given 
already; the deficient weight of the left hemisphere was due to 
a general imperfection in its development, and to the presence 
of a small Flocculus: on the right side the Flocculus was of 
average siza 

The Carpus DmUatum was oblong, and measured '25 inch in 
its longest diameter, which was from before backwards. 

The laminae were not so numerous as usual, and were un«* 
equal in number on the the two sides. 

Superficial kminae: Right side numbered 72. Left side 
numbered 62. 

Specific graoily. The sp. gr. of the entire encephalon was 
found to be 1038, thus dilSering very little from the sp. gr* 
of healthy brains. The sp. gr. of separate parts was not 
taken. 

Chemical Analysis. Professor Boscoe very kindly under- 
took the analysis of the brain for the purpose of ascertaining 
whether or not it was deficient in phosphoriui. I extract from 
his report the following statements. 

For this examination 36'34 grammes of solid brain, plus 
27*20 cubic centimetres of the alcohol in which it was pre* 
served, were taken, this being equal to 4*61 f of the total brain^ 
which, as delivered to him, weighed 790 grammes: the analysis 
of these combined materials yielded a percentage of phosphorus 
of -870. 

The fi^sh brain, however, weighed 28 ounces, which is equal 

to 871 grammes, and thus 100 parts of the dry brain were 

equal to 110*2 of fresh brain; so, to obtain a correct result we 

'37 X 100 
must state it in this form, ^^ , which gives a percentage 

of '336 to the fresh brain. At the time of making this analysis. 
Professor Boscoe also ascertained the percentage of phosphorus 
in a perfectly fresh healthy brain, and found it gave '310^ of 
phosphorus; so that, though the entire quantity of phosphorus 
was less in the brain of the idiot than in the brain of the 



^2 MR BRAI)l.EY« 

healthy man, the percentage was actually higher in the former 
than in the latter \ 

Condition of the bones. The state of the bones was so re- 
markable, that a month after death a rib, unbroken, which had 
been kept in maceration for some time, was removed for ex- 
amination. It was dark in colour, oflfensive in odour, and 
greasy to the touch. Pressure caused the exudation of a thin 
foetid fluid from the ends. The bone could be snapped with« 
very slight forca A scraping placed beneath the microscope 
revealed the presence of vast numbers of oil globules, and gra- 
nules. A thin section at first showed little else; but after the 
oil globules were removed, a process involving much time and 
trouble, the bone-structure itself came into view. The most 
noteworthy peculiarity appeared to be the large size of the 
Haversian canals and interspaces, all of which were filled with 
amorphous granular matter. Interstitial absorption had evi- 
dently been going on in the bones, loosening their component 
parts, and producing the singular softness of structure which 
characterised the bones during life. These changes were pro- 
bably identical with those described by Dr E. Ormerod, as 
found by him in the bones of lunatics ^ 

Remarks. This case presents some points of resemblance 
and some of contrast to the case of a deaf mute, described by 
Dr Broadbent in the November number of the Journal of 
Anatomy and Physiology for 1870, the chief of which appear 
to be the following. 

In Dr Broadbent's case the third left frontal convolution was 
small ; in my case it was well developed. In reference to this 
point it should be remembered that Dr Broadbent's patient was 
dumb during life, whereas the idiot, whose brain I have de- 
scribed, could articulate several words, but seemed to fail to 
grasp their meaning. In Dr Brcadbent's case the annectent 
convolutions were deficient on both sides, so that the parietal 
lobes were almost completely separated from the occipital lobes. 

1 I am indebted not only to Professor Boscoe, bat also to bis laboratory 
assistant Mr BochoU, for tbe time and trouble expended in this analysis. 

* Observations on a peculiar condition of the bones of two insane patients 
. who had fractured ribs. By Edward Latham Ormerod, M.D. Barthol. Hospital 
JteporU, 1870, page 65. 



BESCKIPTION OF THE BRAIN OF AN IDIOT. 73 

In my case this condition existed on one side only, the right, 
where indeed the separation was almost absolute. On the 
left aide, on the contrary, the annectent convolutions were well 
developed. Dr Broadbent also noted that the angular gyrus 
was very small, which was not so in my case. In both cases it 
is interesting to observe that the lobule of the supnirmarginal 
convolution was present on both sides. 

In addition to these points there remain to be noted in my 
case, the existence of a very small and simple Island of ReU, 
extremely ill-developed Corpora Quadrigemina, an almost rudi- 
mentary condition of the vermiform process of the cerebellum, 
and a striking disproportion between the two cerebellar hemi* 
spheres;. 

What weight ought to be attached to the condition of the 
central lobe we probably have not yet the power to decide. 
The small size of the Corpora Quadrigemina is interesting 
when considered in connection with the state of congenital 
blindness which existed. The cerebellar peculiarities were very 
striking, but where the whole psychical and physical states 
were so much jangled and out of tune, it becomes difficult to 
determine to what extent the cerebellum operated as a separate 
pathological factor in the case, whether we regard the idee* 
motor or excito-motor actions of the brain. 

In considering such cases as these the causes of idiocy may 
be conveniently ranged under three heads : 1st, those depend* 
ent upon smallness of the brain itself, which would include 
not only micro-cephalic brains, but all encephala where any 
single part is ill-developed, or rudimentary; 2nd, those caused 
by imperfect anatomical connection of parts; and 3rd, those 
which are the result of imperfection in the cerebral structure 
itself, which would include all those cases where the micro- 
scope reveals imperfect histological development either in the 
cell or fibre elements; and also those cases where, though the 
histological structure appear normal, the chemist discovera 
noteable deficiencies in important cerebral constituents. In 
the case under consideration more than one of these causes 
seems to have been in operation; indeed it is probable that 
all three causes were at work to a certain degree. Let us 
briefly review some of the chief points of interest in this case. 



74 ME BRADLEY. 

The first to notice Is the absence of any traceable predisposing 
or exciting cause; the parents were intelligent, they were not 
related, they could give no history of any case of mental aberra- 
tion, or nervous affection in any shape as existing in the family. 

The bodily condition was interesting; the extreme muscular 
debility, the diseased state of the bones, the general want of 
development in the bodily organs, all would lead one to infer a 
want of structural development, or quality, in the brain-sub- 
stance. It is true that the microscope did not detect anything 
abnormal in the cells or fibres of the grey or white matter, nor 
did the chemist discover any departure from, or want in, the 
most important constituents of the brain ; but, notwithstanding 
all this, it is highly probable that some qualitative excellence 
was wanting; it is at least certain that the non-detection of 
such abnormality proves nothing — ^what microsi^opist or what 
chemist, for example, could describe the nature of the difference 
between two muscles, the one of which was in tone, and the 
other out of tone, other things at the same time remaining 
equal? It is impossible indeed to avoid the impression that 
the want of qucUity, traceable throughout the body, existed also 
in the brain, although we are not able to say precisely in what 
the quality was wanting*. 

Another point necessary to dwell upon for a moment is the 
proportion which existed between different parts of the entire 
encephalon. In spite of the deficiencies in certain parts of the 
cerebellum, it has been seen that it'bore a higher proportion 
to the cerebnmi than is the case in healthy brains; it was not 
absolutely heavier (indeed it was notably lighter) than ordi- 
nary cerebella, but still it was proportionately heavier than it 
is in healthy brains. This is probably to be explained by at- 
tributing a want of normal development, during the first year 
or two of infancy, to the frontal lobes, while the occipital lobes 
and cerebellum continued to grow up to puberty; the character 
of the brain bears out this hypothesis, for although the frontal 
lobes Were not, per se, notably ill-developed, it was evident, 

^ Some interesting statistical tables of the relation between the mental and 
bodily conditions in idiots and the insane is to be found in Dr Gaggenbohl'a 
work, Die Heilung und VerkUtung det Oretinumus, 1853; and in Dr Howe's 
supplement to the Report of the Massachiuetts ComnUesUmert into the eondUum 
of the Idioti of the Commonwealth^ 1868. 



BEdCBIPTION OF THE BRAIN OF AN IDIOT. 75 

from the forward position of the fissure of Rolando, that they 
had not developed pari passu with the hinder parts of the 
brain. The external appearance of the skull too conveyed this 
idea during life; for, while the forehead was very small and 
childlike, the posterior portion of the head was unusually 
wide. The last feature of importance to be noticed is the ana- 
tomical arrangement by which the right occipital lobe was 
almost completely separated from the parietal lobe. I do not 
desire to foist any physiological theorem on to this anatomical 
peculiarity, I only wish to draw attention to it It will be for 
future observation to determine how far such a severance is pro- 
ductive of a want of harmony between volition and locomotion, 
or indeed between volition and action of any kind. 



• 1 

■t 



CONTRIBUTION TO THE ANATOMY OF THE SHOUL- 
DER OF BIRDS. By J. Young, M.D., Professor ofMor 
tural History in the University, Glasgow, 

The shoulder-girdle of the bird has two articulations more 
than occur in the mammal, namely the coraco-scapular and co- 
raco-stemal. While therefore the humerus has its movements 
more restricted in the bird than in man, these two articulations 
increase largely the variations of the angle which the plane of 
the wing makes with that of the trunk. 

The movements at the coraco-stemal joint are antero-pos* 
terior and lateral : they are efifected by special muscles, as well 
as by those which pass from the sternum, or coracoid, to the 
arm or ribs. The special muscles are lodged in the depression, 
on the surface of the anterior lateral process of the sternum. 
The external muscle arises from the greater part of the boundary 
of the depression and the adjacent surface, and is attached to 
the posterior margin of the coracoid. The internal muscle 
arises from the anterior margin of the sternum and the adja- 
cent surface of the depression, and is attached to the upper &ce 
of the coracoid in its posterior half. More or fewer of the fibres 
of the internal muscle pass to the tendon of the external, but 
the origin of the two are always distinct, and the planes of their 
fibres are dissimilar ; so that even when so intimately united as 
in the Green .Woodpecker and the great Goatsucker, this di- 
vergence is very distinct. The action of these muscles is to 
divaricate the coracoids, and to depress the head of these bones, 
thereby rendering the axis of the glenoid cavity more vertical 
llie internal muscle reaches as far as the point of attachment of 
the long slip, sometimes found as a separate part of the coraco^ 
hrachialis brevis, and the fibres sometimes intermingle; but even 
in the Carrion Crow, where such blending frequently occurs, it 
is not found in every specimen, nor even on both sides of the 
same bird. Nor has the coraco-stemalis intemtis anything to 
do with the coraco-brachiaiis longtts, whose origin is entirely 



ANATOMY 6F THE ^SHOULDEB OF BIRDS. V7 

from the inferior face of the hone, whether firom the coracoid 
idone, as in the Oriole, or from that and the sternum, as in 
the Swan, in which there is only one muscular belly, while in 
Larus iridaetylua the sternal and coracoid portions only unite 
near their insertion. It is further frequently said, that the 
cora4X>'8temaiis {extemus) arises from the anterior sternal ribs : 
this I have found only in the Swan, and even there it amounts 
only to a partial second origin. There is in fact confusion be- 
tween this secondary origin and a very distinct muscle, the 
equivalent, on the outside of the ribs, of the triangularis stemi, 
on the inside. Of this external muscle the first slip — that 
from the top of the lateral process of the sternum to the angle 
of the first rib — is always present, more or less recognisable 
by the different direction of the fibres; those of the inner 
muscle passing to the junction of the sternal and vertebral ribs, 
those of the outer not passing so far. Where the external tri- 
angularis is well developed, as in Larus, the bundles pass to 
four ribs; in the Swan only to two. In a young Larus the 
first slip was interrupted by a thin rod of bone, the represen- 
tative of a sternal rib, which was loosely attached to the lateral 
process of the sternum : in the great Goatsucker a tendinous 
slip occupies a corresponding position. This external triar^gii* 
laris is not constant, and its absence, as in the Lapwing and 
Woodpecker, is associated with a high development of the ex- 
ternal oblique muscle of the abdomen. 

The structure of the pectoralis major, the principal rotator 
of the humerus, is such that the movement, necessarily a labo- 
rious one, is performed, so to speak, by relays of muscle, and the 
lafit part of the movement is even more energetic than the first. 
Thus, in the Heron, the anterior third overlaps considerably the 
middle portion of the muscle along the sternal crest, and the 
tendon which lies between the middle and posterior divisions 
affords attachment to the fibres of the latter, so that each por- 
tion provides a fixed point on which the following one may act. 
The fibres which arise from the posterior part of the sternal 
gnrface do not converge to the humerus, but spread out on the 
under surface of the muscle. By this arrangement, apart from 
the dynamical advantages, the nerve is protected from pressure; 



79 fJtowasoH TOinva 

amee tbe more enesgetie the contraction, the laiger is the open* 
ing by whieb it enters the deep face of the muscle. The 
antagonistic rotat<»:, the pectoraUs fnsdiua or suboUivius, gains 
great -power by the spiral which it describes firom its origin to 
its humeral attachment 

But this rotation is only guided by museles. The humerus 
itself is so constructed that rotation is a necessity, and that, 
further, the rotatory movement yields, and can yield, no other 
result Ihan such inclination of the wing to the horizon as 
Pettigrew has found to be the normal one in 6jght. Marey» 
in working out the figure of eight, which Pettigrew had pre* 
viously dem^mstrated to be the typical form in ihe muscular 
arrangement of hollow organs, says that the form of the 
«houlder-joint permits of the appropriate movement, which he 
everywhere treats as the work of the muscle& Now the form of 
the humeral articular surface is such that the pull of the great 
pectoral is much greater than would be required if rotation 
of the axis of the humerus were alone necessary. The sur* 
face is divisible into three area^: a ridge (if X may use such 
a word for the line of juncti<m of two dissinuls^ curves) 
passes from the upper axid front part of the {^ticular head, 
obliquely downwards and inwards, so as to form an angle of 
Wfi with the long axis of the bead, and of the condyles at tbe 
inferior end of the bone. To the outer side hes a helicoid 
surface, which dips very suddenly into the bicipital groove. 
To the inner side, the upper two-thirds of the articular surface 
is the segment of a very large circle, the plane of whose dia* 
meter is at right angles to that of the bone : the inferior third 
is a small segment of a hoge circle, whose diameter plane is at 
45^ to that of the bone ; this surface likewise dips abruptly on 
to the general surface of the bone. The helicoid surface travel 
on the coraco-scapukff ligament by what I have called the ridge, 
and the rotation of the humerus involves the partiaji pro* 
niiticai of the limb, without muscular aid at all. In flexion, 
contrariwise, the movement is reversed, apart from muscular 
puU. But both peotoraUe major and p. medium act on the long 
handles of the levers, the salient crests : both have to deal with 
rotationi but the size of the laiger muscle has reference, besidea 



ANAT0M7 m THE SBOUUXKB, OF 6IBDS. 79 

oiher tbizigs, to ihe curve which the shaft of the bone makes 
in the air. To depress the bone when the shaft is at the 
summit of its cmre is the duty of the most posterior fibres of 
the musda The p, medius^ on the other hand, is too small to 
act as an efficient elevator of the limb, but its size is quite 
sufficient, oonddering the advantage which its spiral course gives 
it, to undo the curve described by the humerus in the forward 
and downward stroke of flight. 

Martius» in treating of the torsion of the humerus, says;, 
that in man it has travelled through 180^ in birds 90^ But if the 
aids of the articular sur&ce is identical in both bones, relatively 
to the axis of the condyles, the torsion must be equal The ao* 
oompanying figures were drawn for the purpose of showing that 
the ridge of the bird's humerus occurs also in man. The diffi- 
culty of representing on a flat surface the details of the curves, is 
only partly overcome by the nature printing in the accompanying 
figures. The head of the humerus was sliced at equal distances, 
parallel to, and at right angles to, the axis of the condyles, 
and the outlines of the slices traced on paper. From the com- 
parison of these figures it becomes evident that the rounded sur- 
fiEbce is not hemispherical, but is made up of the segments of two 
circles, that with the larger radius being on the side of flexion 
(the anterior in man), that with the smaller, on the side of exten- 
sion (or posterior), and that the line of junction of these two is 
not in a vertical line, but slightly inclined from above, and 
in front downwards and backwards, in the same relative direc- 
tioB as in the bird, and at 25^ to 30^ inclination to the axis of 
the condylea The steep faces, which in the bird give to the 
dose of flexion and extension something like a jerk, are not 
found in man ; but the spiral movement i& distinct enough in 
the pronation of the hand, when the limb is laid across the 
chest. Thus even in the details of a single joint the unity 
of plan of the vertebrates is curiously iUustrated, and the de- 
viation from that plan asserted by Martins appears not to exist, 
the torsion in mammals, reptiles, and birds being alike. 

The movements of the glenoid cavity I have not yet ex- 
amined experimentally with sufficient cara The rarity of allu- 
flioDS to the mobility of the joint is to be explained by the study 



80 PROFESSOR TOUNa. 

.t>f skeletons, in "which the amount of mohility is no longer 
recognisable. But in the Goatsucker I have found that the 
extreme possible divergence of the scapula on the coracoid is 
2Vy the average angle from the centre of the glenoid to the tip 
of the scapula on one side to the centre of the posterior sternal 
margin on the other being in the dried skeleton 57^. In 
the Cinnyris, one of the Nectarinia, the limit of movement ia 
only 8^ that is 47" as the average, 55° as the extreme. Though 
my materials are incomplete, it may be worth while to state 
some of the average measurements of the angle between coraco- 
sternum and scapula, it being borne in mind that the variations 
«re due to the elevation or depression of the sternum, the sea* 
pula being very nearly fixed in its relations to the vertebral 
<x>lumn. 

Bateatrica pavonina 42°, Lestris Richardsoni 43°, 

Pelican 65^ Halisetus aJbicilla 80^ 

Macau 52°, Alca torda 85*. 

From specimens stripped of their muscles it would be un- 
safe to calculate the extreme movement, but the statement of the 
great range of averages — 35® — 80°, may induce some one with 
better opportunities than I possess to carry on the investigation. 
Those who deUght in special adaptations- have in the humerus 
an imperfect example. There are other joints showing still 
more remarkable mechanical results, and with these I am at 
present engaged. The anatomical data for reasonings regarding 
the mechanism of progression have acquired fresh interest of 
late years, and it is only just for me to acknowledge the assist* 
ance which Dr Pettigrew, by his published papers and oral 
communications, has rendered me in studying a subject which 
in this country he has made peculiarly his own. 



AKATOMT OF THE SHOULDEB OF BIBDS. 



81 




Ulnar. 



BadiaL 



Frofilea of transyerae Tertioal aliees of head of Hmnerus (human). L Beetion 
next abduotion Bide of bone ; Y. that on adduction siae. 




Profiles of longitadinal Tertioal fllioes. I. Slioe next radial aide ; 

lY. that next ulnar. 



VOL. VI. 



6 



CONTRIBUTIONS TO THE ANATOMY OF THE IN- 
DIAN ELEPHANT\ Part L, THE THORACIC VIS- 
CERA. By Morrison Watson, MD., Demonstrator of 
Anatomy, University of Edinburgh^. Plate VL 

Owing to the kindness of Professor Turner, I have recently 
had an opportunity of dissecting the thoracic viscera of an In- 
dian Elephant {Elephaa Indicus), and the results of this ex- 
amination I now venture to make public, not so much with 
the idea that I shall be able to communicate much that is al- 
together new, but rather, by adding to the descriptions already 
published, which differ in many points from one another, that 
of the individual which I have examined, and so clear up some 
points in the anatomy of an animal which we too seldom have 
an opportunity of dissecting. At the same time I hope to be 
able to call attention to certain arrangements, which, so far as 
I am aware, have, up to the present time escaped observation, 
and which may therefore prove not altogether uninteresting. 

The thoracic portion of the digestive, circulatoiy, and req)i- 
ratory oi^gans will be considered consecutively. 

Digestive. 

It has long been known that both species of elephant 
have the power of withdrawing water, stored within the cavi- 
ties of the stomach, by means of the trunk inserted into 
the mouth. Sir Emerson Tennent says': "I have elsewhere 
described the occurrence to which I was myself a witness, 
of elephants inserting their proboscis in their mouths, and 
withdrawing gallons of water, which could only have been con- 
tained in the receptacle figured by Camper and Home, and of 
which the true uses were discerned by the clear intellect of 
Professor Owen." I was not, till very recently, aware that a 

1 The viscera of this elephant formed part of the ooUection of the late Pro- 
fessor Goodsir, purchased after his decease by the University. The animal waa 
bought from a travelling menagerie by Mr Goodsir in 1856. — ^W. T. 

* An abstract of this Paper was read before Section D at the Edinburgh 
Meeting of the British Association for the Advancement of Science, AugUBt. 
1871. 

• Ceylon, ii, 816. 



ANATOMT OF THE INDIAN ELEPHANT. 83 

similar observation as to the remarkable habit of the elephant, 
had been made by the author of the Ayeen Akberry, in his ac- 
count of the Feel Kaneh, or elephant stables of the Emperor 
Akbar, in which he says: "An elephant frequently with his 
trunk takes water out of his stomach and sprinkles himself 
with it, and it is not in the least ofifensive/* The means by 
which this is effected Tennent believed to lie in the presence of 
a muscle described by the late Prof. Harrison of Dublin, which 
is referred to as follows in the work already quoted*. Dr Har- 
rison in the course of his examination of the thoracic viscera, 
observed that an unusually close connection existed between the 
trachea and oesophagus, which he found to depend on a muscle 
unnoticed by any previous anatomist, connecting the back of 
the former with the fore part of the latter, along which the 
fibres descend, and can be distinctly traced to the cardiac orifice 
of the stomach. Imperfectly acquainted with the habits and 
functions of the elephant in a state of nature, Dr Harrison 
found it difficult to pronounce as to the use of this very peculiar 
structure; but looking to the intimate connection between the 
mechanism concerned in the functions of respiration and deglu- 
tition, and seeing that the proboscis served in a double capacity 
as an instrument of voice and an organ for the prehension of 
food, he ventured to express the opinion that this muscle, 
viewing its attachment to the trachea, might either have some 
influence in raising the diaphragm, and thereby assisting in 
expiration, "w that it might raise the cardiac orifice of the 
etofnach, and so aid this orgcm to regurgitate a portion of its 
contents into the oesophagus^r Unfortunately, however, for this 
theory, no trace whatever of the trachea-oesophageal muscle was 
to be found in the specimen I examined ; and, as no mention is 
made of it by any other anatomist, we must, I think, conclude 
that the muscle was only exceptumaUy present in Harrison's 
elephant (which is also the case with a corresponding muscle in 
the human subject"), as it is hardly probable, supposing the 
function alluded to to depend exclusively on the existence of 
such an arrangement, that the muscle should be present in one 
specimen and absent in another. 



* lb. n. 814. « Proc, Roy, IrUh Acad, iv. 183. 

* Vide Henle, AnatomU des Mensehefif Band n. p. 151. 



6—2 



84 MR WATSON. 

The muscular fibres of the oesophagus are distinctly striated 
even down to the oesophageal opening in the diaphragm, and 
are arranged in two layers, an external, the fibres of which 
are distinctly longitudinal in direction, and an internal, which 
consists of two sets of spiral fibres, one of which passes firom 
right to left, whilst the other passes in the opposite direction, 
and thus gives rise to a decussation of the fibres at all points. 
It is thus to be observed that the arrangement of the fibres 
differs materially firom that described by Prof. Rutherford in 
the gullet of the ruminant\ 

With reference, therefore, to the regurgitation of water, we 
must, I think, conclude that, in the absence of any peculiarity 
in the stomach to account for it» in all probability it depends, 
as Flourens has shown to be the case, in the physiological re- 
gurgitation of the ruminant, and Magendie in the pathological 
regurgitation of other animals, on the action of the diaphragm, 
and abdominal muscles. 

Thoracic duct. This vessel presents the usual arrangement 
as far as its relations are concerned. At the posterior extremity 
of the thorax it has a diameter equal to that of a goose-quill, 
whilst towards its termination it has re&ched the size of the 
axillary artery of the human subject. No appearance of the 
reservoir of Pecquet, which is described in the Encydop^die 
M^thodique\ as being placed opposite the third dorsal verte- 
bra, and extending for a distance of several inches was to be 
seen, the canal being of uniform diameter throughout A single 
pair of valves was placed about two inches behind the spot 
where the vena azygos turns downwaxds to open into the right 
anterior cava, the presence of such being altogether denied in 
the article just quoted. 

Organs of Circulation. 

PericardiunL Fig. 1. The description of this viscus will 
be better understood after a few words concerning the dia- 
phragm. As this muscle had been removed along with the 
viscera from the cavity of the thorax, and consequently coxi- 

* Journal of Linnean Society, vni. 

* Encyelopidie Mithodique, Article Paohydermes. 



ANATOMY OF THE INDIAN ELEPHANT. 85 

siderably injured before I commenced the dissection, the descrip- 
tion is not so perfect as might otherwise have been the casa 

The greater portion of the diaphragm was occupied by the 
central tendon, from the margin of which the muscular fibres 
passed off to their attachment to the circumference of the tho- 
rax. These two parts differed materially from one another, for 
whilst the central tendon was very thick and strong, the mus- 
cular portion of the septum, on the other hand, was thin, and 
the bundles easily separated. An arrangement, however, to 
strengthen this deficiency was present, consisting of two plates 
of yellow elastic tissue, one covering each of the surfaces of 
the muscular portion of the diaphragm. Both of these plates 
of elastic tissue passed imperceptibly into the substance of the 
central tendon and there lost themselves. The anterior plate 
differed from the posterior in this, — that, whilst the posterior 
plate covered every part of the muscular portion of the dia- 
phragm, the anterior was confined to that part of it which 
came into relation with the base of the pericardium, and 
shaded off beyond this region to the left side into a muscular 
fascia of considerable strength, and composed of fibrous tissue, 
whilst to the right, owing to the diaphragm having been cut, 
I could not follow it to its termination. This much however 
is to be said, that a considerable admixture of fibrous with 
the yellow elastic tissue having already taken place, seemed to 
imply that on the right, as on the left side, this elastic plate 
gradually shaded off into a fibrous tissue. If this supposition 
be correct, then we must regard the anterior elastic plate as 
being connected physiologically with the pericardium, and not 
with the diaphragm. The probability of this view will be in- 
creased when the attachment of the pericardium to the dia- 
phragm is considered. 

John Hunter* states that " The pericardium adheres to the 
diaphragm nearly as in the human, but not so closely and 
firmly;" but a more correct description would be that it forms 
an intermediate stage between the completely free pericardium 
as described by Prof. Turner in. the walrus*, and the closely ad- 
herent human pericardium. Its posterior surface is smooth, 

^ Euayt and Obtervations by Owen, ii. 172. 

• Journal of Anatomy and Fhytiology, November, 1870. 



86 MB WATSON. 

and rests against the anterior surface of the tendinous portion 
of the diaphragm. It is not, however, attached to this as in 
man; but the corresponding surfaces of the diaphragm and pe- 
ricardium being smooth, evidently permit a considerable amount 
of motion between the opposed surfaces. This motion, however, 
must be limited by the presence of two stout bands of yellow 
elastic tissue, which become continuous with the fibrous sub- 
stance of the pericardium at its posterior extremity. These 
two bands are flattened from above downwards, and measure 
each about an inch in breadth. They are separated from one 
another at their attachment to the pericardium by an interval 
of about three inches. From this attachment they diverge 
from one another, passing backwards and downwards and finally 
become continuous with, and lost in the elastic plate already 
described as covering a portion of the anterior surface of the 
diaphragm. It is a fact worthy of notice that the pericardiac 
band of the right side was entirely composed of that peculiar 
striated form of yellow elastic tissue which, so far as I am 
aware, has only once before been described, and that in the 
ligamentum nuchae of the giraffe by Mr Quekett\ In the left 
band, as well as in the plate into which the bands expanded, 
the elastic tissue presented the usual appearance^ This mode 
of attachment of the pericardium to the diaphragm in the ele- 
phant seems up to this time to have been overlooked, the near- 
est approach to an accurate description of it having been made 
by MM. Vulpian and Philipeaux*, who say: "Au sommet du 
coeur le pc^ricarde so termine en pointe obtuse, et se continue 
directement en ce point avec un fort cordon fibreux qui par 
son autre extre^mit^ va s'ins^rer au centre phr^nique du dia- 
phragme." This description, however, is incomplete, though 
nearer the truth than those given by other anatomists. As re- 
gards the anterior attachment of the pericardium nothing need 
be said, farther than that it becomes continuous with the outer 
coat of the ascending and transverse parts of the arch of the 
aorta. 

Heart, With regard to the external configuration of this 
organ the reader may be referred to the very excellent descrip- 

* Histological Catalogue^ i. 89. 

• Ann, des Sc, Nat, Zoologie, 1856, Vol. v. 



ANATOHT OF THE INDIAN ELEPHANT. 87 

tion of MM. Yulpian and Philipeaux already noticed. The 
only point to which they have omitted to give prominence, 
being the distinct bifidity of the apex of the organ, a fact 
which corroborated the statement of iElian, Stukeley^ and 
Mayer*. This bifidity of the heart is interesting when taken 
in connection with the singular and much more pronounced 
separation of the ventricles in the dugong and manatee, and its 
occasional occurrence in the porpoise. No trace of an os cordis 
was to be found in the wall of the heart. The walls of the 
cavities in general are much thinner than one would expect in 
so large an animal; they are however, especially on the right 
side of the heart, extremely elastic. 

On opening into the cavity of the right auricle (fig. 2), five 
distinct openings were seen, viz. those of three vensB cavse, two 
anterior — right and left — and one posterior; that of a single 
coronary vein ; and that of the ventricle. At the mouth of each 
of these openings was placed a valve, with the exception of that 
of the coronary vein, which lay under cover of a pectinate 
muscle. The valve at the mouth of the posterior cava — the 
Eustachian — was of large size, and an-anged in the usual man- 
ner. That at the mouth of the right anterior cava was peculiar. 
Taking its rise from the inner aspect of the opening it wound 
round the upper margin, and finally passing backward became 
attached to the outer wall of the auricle close to the point of 
entrance of the posterior cava. This description precisely cor- 
responds with that given by Vulpian and Philipeaux, with this 
exception, that in their specimen the valve passed round the 
lower margin of the caval opening instead of the upper. It is 
remarkable that their description of the valve corresponds ex- 
actly with that given by Prof Turner" of one which he met 
with as a very rare variation in the human subject — ^a fact not 
without interest to those who believe " that man still bears in 
his bodily frame the indelible stamp of his lowly origin." At 
the mouth of the left anterior cava was situated a small fene- 
strated valve formed by a duplication of the endocardium, which 
is not generally present in those animals possessed of two ante- 

^ E$$ay towardt the Anatomy of the Elephant, Lond. 1723. 

* Nova acta Acad. Ca$. Leo^Car. Vol. xxn. 

' Proe. Boy, 8oc, Edinburgh, and Journal of Anatomy, Sfay, 1S69. 



88 MB WATSON. 

lior vensd cavse^ and which I think we mnst regard as a rudi- 
mentary appearance of the Thebesian Talve of the higher mam- 
mals. Mr Marshall, in his paper on the development of the 
great veins ^ says: ''As to the Thebesian valve it is present in 
every instance in which the left venous trunk forms a coronary 
sinus, receiving veins from the heart alone as in man, the mon- 
key, &c. ; but amongst those animals which have a left azygos 
or left superior cava» it is certainly absent, as in the calf, hog, 
&cJ* The presence of this rudimentary valve in the elephant, 
therefore, is of interest, not only as establishing a fact not pre- 
viously known, but also as showing the accuracy of Mr Marshall's 
observation, that the coronary sinus is the homologue of the 
lower part of the left anterior cava, since, in the case before us, 
we not only have the great coronary vein opening into the left 
anterior vena cava^ and defended by the usual pair of valves, 
but we also have a rudimentary Thebesian valve, which is not 
usually present in those animals in which the entire left duct of 
Cuvier is persistent. The fourth opening was that of the an- 
terior coronary vein, which opened on the right wall of the 
auricle, and, as already stated, was not defended by valves. 
Yulpian and Philipeaux describe a third coronary vein as open- 
ing by a distinct orifice into the auricle, but this was not the 
case in the present specimen. The right auriculo-ventricular 
opening was guarded by the tricuspid valve, which however 
was provided with a small additional cusp placed between the 
cusps which correspond with the anterior and internal cusps of 
human anatomy. 

The cavity of the left auricle (fig. 3) presented the four open- 
ings of the pulmonary veins, two of which were derived from 
each lung. Vulpian and Philipeaux describe only two openings 
in connection with this auricle, a statement difficult to reconcile 
with the dissection of the present specimen. In front of these 
openings was placed a valvular structure, which however (by 
reason of its small size) could not have been effective in pre- 
venting the regurgitation of blood into the veins. It arose from 
the anterior portion of the auricle, and passing down in front of 
the openings was inserted posteriorly into the anterior raised 
margin of the fossa ovalis, which was well defined on this, as on 

» Phil Tram. Part 1, 1860. 



ANATOMT OF THE IKDIAN ELEPHANT. 89 

the right side, by several tendinous cords. The relation of this 
Yalve to the openings of tbe pubnonary veins was similar to 
that of the valve of the right superior cava to its opening. This 
valve does not seem to have been previously recognised. The 
mitral valve was composed of three distinct cusps, the third 
small cusp being situated between the inner extremities of the 
cusps which correspond with the larger anterior and posterior 
cusps of human anatomy. 

The ventricular cavities presented nothing remarkable, and 
I may refer those desirous of having a full description of them 
to the memoir by MM. Yulpian and Philipeaux already 
quoted. 

Puhnonary artery. This vessel entirely concealed the as- 
cending part of the arch of the aorta from below, and presented 
three well marked dilatations externally, which corresponded to 
the position of the sinuses of Valsalva. Passing forwards and to 
the right it divided in the concavity of the aortic arch into its 
two branches, one of which passed to each lung. The ductus 
arteriosus was attached not to the left branch of the pulmonary 
artery, as in the human subject, but to the trunk of that vessel, 
three inches behind its point of bifurcation. In front it was at- 
tached to the concavity of the aortic arch, just beyond the place 
of origin of the left subclavian artery. The small azygos lobe 
of the right lung did not receive any separate branch from the 
pulmonary artery before it entered the substance of that lung. 

Pulmonary veins. These were two in number on each side; 
the anterior of which was formed close to the hilum pulmonis 
by the union of several branches derived from the apical lobes 
of the corresponding lung; whilst the posterior was derived from 
the posterior or basal part of the lung of its own side; but the 
vein of the right side received in addition a branch of consider- 
able size from the azygos lobe of this lung, which opened into it 
immediately before it entered the auricle. These four veins all 
opened by distinct orifices into tbe left auricle close to one 
another. 

Coronary arteries. According to Camper* there is but a 
single coronary artery, which shortly after bifurcates so as to 
supply the usual branches to the heart. In the present dissec- 

* Description anatomique cfun 6Uphant mdU. 



90 MB WATSON. 

tion, as in those of Vulpian and Mayer, there were two arteries, 
which were distributed in all respects in the usual manner to 
the substance of the heart. 

, With regard to the number of branches given off from the 
arch of the aorta there is a difference of statement. For whilst 
Cuvier* and Mayer' assert that there are three trunks given off, 
viz. the right subclavian, a trunk common to the two carotids, 
and the left subclavian: on the other hand, Hunter', Tiedemann, 
and Vulpian ^ only mention two, viz. 1st, the innominate, which 
gives off the right subclavian, and the two carotids; and 2nd, 
the left subclavian. In the present dissection the arrangement 
was in accordance with the statement of the authors last men- 
tioned (fig. 1). In addition, however, the trunk of the inferior 
thyroid artery was given off from the point of sepai'ation of the 
two common carotids, a point which Mayer was the first to notice, 
although, from the fact of Camper having omitted to mention 
it, he seems doubtful whether it was not to be set down as a 
variation in his specimen. None of the other authors quoted 
make mention of this arteiy. This point is interesting as show- 
ing in one of the lower animals a condition which only occurs 
exceptionally in the human subject, that is, the presence of 
an arteria thyroidea ima coming off from the trunk of the 
innominate. 

Anteiior vence cavas. — Each was formed by the junction of 
ihree large trunks a short distance in front of the arch of the 
subclavian artery. Of these, one came from the outside, a 
second came from the direction of the middle line, whilst an in- 
termediate one passed directly backwards. The vena cava of 
each side, thus formed, passed directly backwards, receiving in 
its course several smaller veins, one of which was the trunk 
formed by the union of the companion veins of the mammary 
artery, and finally opened into the right auricle. In addition 
to these the right anterior cava received the azygos vein imme- 
diately before piercing the pericardium. There was no trace of 
a small or left azygos vein: the posterior cava immediately 
after piercing the diaphragm opened into the auricle. 

^ Lemons d'afuttomie eomparSe, Paris, 1802. 
' Nov, Act. Acad, Cat, Leo-Car. Vol. xxn. 
* Eatay$ and Observatunu by Owen. * Ann. Sc, Nat. YoL v. 



Al^ATOMT OF THE INDIAN ELEPHANT. 91 

Coronary veins. — These are two in number, a great, and 
small. The great coronary vein commenced toward the apex 
of the heart by the union of several branches, the trunk formed 
by which lay in the anterior ventricular groove. It then turned 
to the left along the left auriculo-ventricular groove, received in 
this course a large vein from the posterior- ventricular groove, 
and opened finally into the left anterior vena cava, a short dis* 
tance from its termination in the auricle. A pair of valves 
guarded the entrance into the cava. 

The small coronary vein, corresponding to the small anterior 
vein of Galen of human anatomy, commenced by the imion of a 
number of small twigs toward the left side of the base of the 
right ventricle. It then passed from left to right along the 
right auriculo-ventricular groove, and opened into the right 
auricle by a distinct opening which was not guarded by any 
valve. Vulpian and Philipeaux described a third coronary vein 
as opening by a distinct orifice into the auricle, but such was 
not to be found in the present dissection. 

Respiratory System. 

Trachea. This tube, so far as it lay in the cavity of the 
thorax, consisted of twelve rings, which were aU incomplete 
behind. These rings presented a peculiarity which, so Ceix as I 
am aware, has not been observed in the trachea of any other 
animal ; that is, the subdivision of the different rings into dis- 
tinct segments by means of distinct synovial joints. These joints 
were each enclosed in a distinct capsule, and presented all the 
characters of perfect articulations (fig. 4). The number of these 
articulations in any given ring varied, as many as three being 
found in a single ring; in others, only two or one, and in others 
none at all. The same peculiarity manifested itself in the bron- 
chial cartilages. Mr Bishop ^ in his reference to the trachea of the 
elephant, says, '' Trachea thirty rings, which are often partially 
subdivided." Whether this applies to the peculiarity men- 
tioned, or whether it refers only to that incomplete formation 
of the separate rings, so common in many animals, I am at a 
loss to determine, as beyond the statement quoted he makes no 
further reference to the subject. 

^ Cyelop48dia of Anatomy t article Voice. 



92 MR WATSON. 

In addition to and internal to the rings, the trachea pre* 
sented a strong lining of yellow elastic tissue, the fibres of which 
were longitudinal in direction. This lining passed uninter- 
ruptedly from the trachea into the bronchial tubes. Resting 
directly against this elastic plate was the tracheal mucous mem- 
brane^ which was thrown into distinct longitudinal folds or 
rugsB (fig. 5). 

The interval posteriorly between the ends of the rings, in 
addition to the structures just mentioned, was completed by two 
distinct layers of involimtary muscle. One of these, the more 
superficial, passed directly across the interval so as to be at- 
tached to the extremities of the rings of the trachea. The 
deeper set arose from the V-shaped projection of the last tra- 
cheal ring, the central fibres passing vertically upwards beneath 
the transverse fibres of the superficial layer, the lateral bundles 
assuming a more and more transverse direction, so that the 
most external of these formed the commencement of the series 
which filled up the interval between the extremities of the 
bronchial cartilages. 

Bronchi, The right bronchus consisted of eight, the left of 
six rings. In addition to these, however, there were several 
small cartilaginous nodules of small size representing rudi- 
mentary rings. No accessory bronchus such as is so common 
among the ruminants was met with; the azygos lobe of the 
right lung receiving its air-tube fi-om the right bronchus after it 
had entered the substance of the lung. 

Lungs, Each of these presented an elongated oval form, 
the length being about twice the breadth. Only two surfaces 
are to be distinguished in each, a dorsal and a ventral; or, 
more properly speaking, an external and internal. There 
was thus no distinct diaphragmatic surface to be recognised, 
as in the human subject; the base of the lung, on account 
of the peculiarly elongated condition of the thorax dorsally, 
being continuous with the ventral surface, and being wedged 
in, as it were, between the diaphragm and the upper wall of the 
thorax. The line of junction of the two surfaces thus gave rise 
to the margin of the lung, which was marked externally with 
distinct marginal notches similar to those described by Oweu^ 

* Anatomy of the Vertebrates^ VoL ui. 



ANATOBCT OF THE INDIAN ELEPHANT. 98 

in the lung of the Khinocero& The hilum of the lung was 
situated on its internal surface at the junction of the anterior 
with the middle third o! its length. The substance of each 
lung was divided into a number of lobes by processes of a 
lax connective tissue, which could be easily torn so as to 
separate the lobes. These rudimentary lobes were much more 
numerous towards the apex than at the opposite extremity 
of the organ, and they corresponded exactly in nimiber and 
form in the lung of each side. The only difference in the 
two lungs consisted in the possession by the right of a small 
quadrangular azygos lobe which, connected to the organ iia- 
mediately below the hilum, projected transversely towards the 
left, lying between the vena cava inferior and the anterior 
surface of the diaphragm. Yulpian and Philipeaux describe 
three distinct lobes in the right lung, and none in the left 
This^ however, was certainly not the case in the present disseo- 
tion. The arrangement of the pleural sacs could not be deter- 
mined on account of their lacerated condition. This membrane, 
however, did not separate the lobes from one another, but 
passed continuously over the surface of the lung. The pleura 
were devoid of all trace of yellow elastic tissue ; but upon strip- 
ping them off their respective lungs, each of these last was 
found to be be invested as fax forward as the hilum with a thick 
covering of yeUow elastic tissue, which gradually thinned off 
towards the apex, although it could be traced as a separable 
membrane over the entire surface of the lung. This invest- 
ment is much thinner on the internal than on the external 
sor&ce. This covering towards the posterior extremity of the 
lung was almost entirely composed of yellow elastic tissue, 
although containing a slight admixture of ordinary fibrous tissue, 
but on passing forward to the front of the lung, the quantity 
of the latter increased so that at the apex the membrane now 
became very thin so as to be almost entirely composed of it. 

The bronchi on entering their respective lungs branched into 
a number of subdivisions, all of which passed to the apical lobes, 
with a single exception. This, which was the largest of all, 
passed backwards in order to supply the larger posterior portion 
of the lung, giving off in its course numerous branches for the 
supply of the entire organ. The mode of subdivision of the 



94 MB WATSON. ANATOMT OF THE INDIAN ELEPHANT. 

bronchi, however, was not dichotomous, bnt quite irregular. As 
regards the structure of the bronchi themselves, upon entering 
the lung they at once lost all trace of cartilaginous rings, no 
trace of this tissue being found in any part of their ramifica- 
tions. Throughout their entire extent the mucous membrane 
was thrown into folds as already described in the trachea. 

This peculiar non-cartilaginous condition of the intra-pul- 
monic portion of the bronchi has been recognised before in the 
howling monkey^ ichneumon', and several other animals, but 
not, so far as I am aware, has it been previously noticed in the 
elephant 

Upon transverse section under the microscope the bronchi 
were found to be composed from without inwards, of, first, 
a layer of strong elastic tissue; second, a circular layer of 
involuntary muscular fibres; thirdly, a layer of elastic tissue, 
but of a finer quality than that composing the external coat; 
and lastly, of the mucous membrane, the minute structure of 
which could not be satisfactorily investigated on aocoxmt of the 
age of the specimen. 



EXPLANATION OP FIGURE ON PLATE VL 

Fig. L Heart, pericardium and diaphragm. The pericardium 
has been out through and partially turned down to shew the great 
vessela a, Left auricle. 6, Bight auride. o, Pulmonary artery. 
df Aorta, e, Innominate artery, giving off the right subclavian. 
/, Right and lefl common carotids; gg, and the inferior thyroid artery 
between the latter. A, Left subclavian artery, n, Right and left 
anterior venie cavn thrown down, k, Periccurdium with the two 
bands kk. I, Diaphragm, m, Anterior elastic lamina. 

Fig. 2. Cavity of right auricle, a, Mouth of right anterior cava. 
bf Mouth of lefb anterior cava with rudimentary valve of Thebesius. 
c, Posterior vena cava, d, Yalve at mouth of right anterior cava, 
e, Eustachian valve. 

Fig. 3. Cavity of left auricle, a, Mouths of pulmonary veins. 
b, Yalve in connection with these. 

Fig. 4. View of trachea from the front, cuta, Synovial articula- 
tions of tracheal rings. 

Fig. 5. View of trachea from behind, shewing rugose condition 
of the mucous membrane (a). 

^ Carier, Lefom tTanatomie eomparie, Tom. nr. p. 811. ' Ibid. p. 818. 



ON THE ACTION OF INORGANIC SUBSTANCES 
WHEN INTRODUCED DIRECTLY INTO THE 
BLOOD. By James Blake, M.D., F.RC.S., San Fran- 
ciscOt CalifomuL 

Is the present paper I propose to relate some ezperiments 
which have been made by introducing the salts of platinum 
and of palladium directly into the blood. The manner in 
which the experiments were conducted was the same as de- 
scribed in former communications (see this JoumcU for May, 
1870). 

Exp. 1. A dog not very well nourished, weight about 14 lbs. ; 
pressure in the arteries 4 to 4*6 inches ; heart's action regular. 
A solution containing 1 gr. of chloride of platinum in 3iij of 
water was injected into the jugular vein ; in 12" the action of 
the heart quicker, oscillations not more than 0*1 to 0*2 inches ; 2(/' 
action of heart irregular, pressure increased to 5*5 inches; 45" 
pressure diminishing; at 2^ it was 2 or 2*2 inches, oscillations 
slight, breathing rather irregular ; 6' inject 2 grs. ; no perceptible 
effect on the heart; in 40" pressure in the arteries down to 1*25 
inches; no expression of pain, animal quiet, although sensation 
apparently perfect; 2^ no oscillation in the mercury, although 
the heart could be felt pulsating; respiration suspended for 50", 
the animal lying quite still, although sensible; no convulsion, 
no expression of pain; pressure in arteries 1*5 inches; slight 
oscillations (about 0*05 inch). After 50" respiration again com- 
menced and continued quite regular, but rather slow for about 
2^; suspended for 1', and again commenced; 5' inject 2 grs.; 
pressure 1 inch; respiration stopped; in 40" heart's action con- 
tinuing after respiration had ceased. On opening the thoi*ax 
the heart was found beating slightly, both cavities contained 
blood, right much distended ; the blood in the right cavities was 
dark; that in the left was brighter, a dark maroon colour, co- 
agulated imperfectly; lungs anaBmic-collapsed. 

Exp. 2. Dog weighed about 16 Iba A solution containing 
2 grs. of chloride of platinum was injected into the axillary 
artery, the pressure being 4 to 5 inches. Immediately violent 



96 MR BLAKE. 

struggles and expression of pain. On the animal becoming 
quiet, the pressure was found to have increased 2 inches; 2' ac- 
tion of heart slower, pressure down to 3"5, respiration suspended; 
for 30" animal sensible, no expression of pain; 5' inject 4grB., 
violent struggles; 45" animal quiet, pressure 9 to 10 inches^ 
heart's action quieter, respiration slow, deep, arrested; at 2^ 
heart's action continues, pressure 5 '6 to 6 inches, one deep re- 
spiratory movement after respiration had been suspended; 1''30" 
no farther sign of life, except that the heart continues beating; 
8' after respiration had ceased, the pressure was still 3 inches; 
.slight oscillations from the action of the heart perceptible. On 
^opening the thorax the heart was found contracting feebly; both 
cavities contained blood, which was dark in the right, rather 
•brighter in left (dark maroon colour). The lungs were much 
congested, and there were several ecchymosed spots immediately 
under the pleura, and in the substance of the lung. 

Exp. 3. Injection of chloride of platinum into the veins. 
GbnersJ symptoms. A strong healthy dog weighing about 
16 lbs. Inject into the jugular vein a solution containing 1 gr« 
of the salt in three drachms of water. No immediate expression 
of pain; 45" staggering with a tendency to fall backwards; 2' 
respiration affected longer and deeper, no expression of pain^ 
sensibility perfect; the change in the character of the respiration 
comes on in paroxysms, lasts about 30", and then the breathing 
becomes quite natural; 4' animal laid down on its side, respira- 
tion suspended for 40", no convulsion, no expression of pain, 
sensibility unimpaired, took notice of sounds after respiration 
had been suspended; 45" respiration again commenced, rather 
blowing at first, but soon became natural. This total suspen- 
sion of the respiratory movements recurred four times during 
the next ten minutes, lasting from 45" to 1' 15", during this 
time the animal lay quite still, no sign of pain, or even uneasi- 
ness, although sensibility apparently remained perfect On one 
occasion, after respiration had been suspended for 1' 15", there 
was a slight stiffening of the muscles of the legs and of the tail, 
but nothing like a convulsion; the action of the heart, as felt 
through the sides of the chest, apparently very weak. Inject 
1*5 gr., breathing arrested in 30"; again commenced after being 
suspended 45", heart pulsating feebly, respiration finally sua- 



ACTION OF IKORQAKIC SUBSTANCES. 97 

pended after 3'. On opening the thorax the heart was found 
pulsating feebly, and continued irritable for about 10'; both 
cavities contained blood, the right dark, the left scarlet; it co* 
agulated imperfectly. Lungs rather red, but collapsed perfectly, 
evidently contained but little blood. Partial contractions of the 
voluntary muscles were observed some minutes after death. 



Scdis of Palladium. 

Exp. 4. Dog about 20 lbs. Pressure in the arteries before 
injecting 5 to 8 inches. Inject \ gr. of chloride of palladium in 
\ oz. of water into the jugular vein. In about 12" action of the 
heart fluttering, pressure diminished, oscillations less; 2' respi- 
ration deeper and longer, pressure 4 to 4*2; 5' pressure 3*5 in. 
inject 1 gr. of the salt; 14" the pressure in the arteries suddenly 
fell to 1*5 inches, and no oscillations perceptible, although the 
heart could be felt beating slightly through the walls of the 
chest; 3' respiration arrested. On opening the thorax there 
were partial movements of the heart, but no regular contrac- 
tion; both cavities contained blood; that in the right side dark» 
in the left rather brighter, dark maroon colour, coagulated 
slowly. Lungs very pale; amemic; spontaneous contractions of 
the voluntary muscles were observed 6 minutes after death. 

Exp. 5. Dog weighed about 17 lbs. Pressure in the arte* 
ries from 4*5 to 6 in. Inject ^gr. chloride of palladium into 
the axiUary artery; 12^' there was general spasm with suspen- 
sion of respiration, then violent struggles causing great oscilla- 
tions in the pressure; 45" on the animal becoming quiet, the 
pressure was found to be increased to 12 in., oscillations, 1 in., 
heart's action quick; but regular respiration almost suspended, 
one complete respiratory movement about eveiy 45". This 
lasted 4' 30", and then respiration finally ceased. During this 
lime the pressure in the arteries gradually diminished; but 1' 
after respiration had ceased it was 6 in., heart beating regularly. 
The heart continued beating regularly for T after respiration 
had ceased; and when its pulsations were no' longer perceptible, 
the pressure in the arteries was still 3*6 in. On opening the 
thorax the heart was found contracting irregularly, both cavi<* 

VOL. VI. 7 



98 HR BLAKK, 

ties full of dark blood, lungs congested with ecchymosed spots ; 
the muscles of the leg, through the artery of which the injec- 
tion was made, continued to contract spontaneously 15' after 
death. 

Exp. 6. Dog weighed about 14 lbs. It was not confined. 
Inject J gr. of chloride of palladium into the jugular vein; no 
expression of pain; no marked symptoms; 2' animal seems dull, 
hangs its head, respiration long and deep; after 10' again inject 
^ gr.; 15'' animal fell backwards, then lay on its side, no con- 
vulsion, no expression of pain ; after 40" again rose and walked 
about, but seemed uncomfortable; 2' again fell down, respira- 
tion very long and slow, passed urine and faces; 4' respiration 
suspended completely for 45", animal lying quite still, as if 
dead, but quite sensible, no convulsion. The same thing occur- 
red four times during the next eight minutes, when the respi- 
ration was finally stopped. On opening the thorax the heart 
was contracting irregularly and slightly; the lungs and the 
blood presented the same appearances as in Exp. 4. 



Salta of Iridium. 

Exp. 7. Owing to the chlorides of iridium being decom- 
posed by water, the double chloride of iridium and ammonia 
was used. 

Dog strong, healthy, weight 16 lbs. Pressure in the arteries 
4 to 5 in. A solution containing about ^ gr. of the salt was 
injected into the jugular vein; 8" the pressure in the arteries 
began to diminish, the heart's action became quicker, oscilla- 
tions in the mercury slight: 20" there was an increase in the 
pressure to 6 in., no expression of pain; the pressure again gra- 
dually diminished; at 4' it was 3*6 in., respiratory movements 
slower and deep: 10' inject 1 gr.: 11" pressure falling rapidly: 
30" pressure 1 in., respiration deep, irregular: 1' 20" pressure 
0-6 in., no oscillation, respiration intermittent : 3' 30" respira- 
tion ceased. On opening the thorax the heart was found con- 
tracting rhythmically, right cavities much distended with dark 
blood, left cavities contained blood not so dark, the blood oo- 



ACTION OF mOBQANIC SUBSTANCES. 99 

agulated imperfectly. The lungs were anaemic, collapsed more 
than usual on the thorax being opened \ 

Ajs r^ards the proximate cause of the effects that follow the 
mixing of these substances with the blood, it is evident that 
even in very small quantities (^ gr. chloride of palladium) they 
exert a marked influence on the passage of the blood through 
the lungs; and in larger doses they completely stop the pulmo- 
nary circulation, by causing contraction of the smaller arteries 
or the capillaries. This is shown by the sudden fall in the pres- 
sure in the arteries immediately after the injection of a full 
dose into the veins, and the presence of scarlet blood in the 
left cavity, when the heart continues beating sometime after 
respiration has been arrested. In smaller quantities they pass 
through the lungs after a few seconds, and an analogous effect 
is then often exerted on the systemic vessels, the pressure in 
the arteries becoming increased. This effect, however, on the 
systemic vessels is much better shown when the salt is injected 
directly into the arteries, the obstacle to the circulation be- 
coming so great, that the pressure of the blood in the arteries 
is more than doubled, or equal to a column of mercury of 
eleven to twelve inches. It is probable that this obstacle to 
the systemic circulation is caused by the substance producing 
directly contraction of the smaller vessels or capillaries. In 
the case of the pulmonary vessels it certainly cannot be due 
to any effect on the va9o-motor nerves, at least as far as the 
central organ is concerned, for it so immediately follows the 
injection, that the salt would not have had time to reach 
the nervous centres. The ansBmic condition of the lungs in 
cases where a full dose has been used, and when the salt has 
not even reached the left side of the heart, shows the local 
action on the blood-vessels. The effusion and ecchymosis in 
these organs when the substance has been introduced into the 
arteries, is owing, undoubtedly, to the great backward pressure 
to which they are subject when the heart is contracting under 
the pressure of a twelve inch column of mercury. The presence 
of scarlet blood in the left cavities when the action of the heart 

1 Owing to my notes containing the acoonnt of other experiments performed 
with the BftltB of iridium, and also with those of asmium, having been destroyed, 
I am unable to famish the data as to the farther action of these sabstances. 

7—2 



100 MB BLAKE. ACTION OF INOBaAKIC SUBSTANCES. 

has continued some time after respiration has ceased, shows how 
complete must be the obstruction to the circulation in the 
lungs. The left cavities are prevented from discharging their 
blood by the overdisturbed state of the right heart*. 

The suspension of the respiratory movements for more than 
a minute without producing any distress, the animal being per- 
fectly sensible, their renewal and continuance for some time, to 
be again suspended, and this occurring many times, is certainly 
a curious phenomenon, and one difficult to reconcile with the 
accepted theories as to the cause of the respiratory movements, 
unless, indeed, the presence of the salt in the blood can, to a 
certain extent, supply the place of oxygen. This hypothesis 
receives support from the fact, that the heart continues beating 
so long after respiration had ceased (7 minutes), and this too 
when the pressure in the arteries was at four inches, or about 
its normal height. The proximate cause of death, when a mo- 
derate quantity of these salts is introduced into the veins, is, 
undoubtedly, owing to arrest of the respiration; but whether 
this is caused by their action on the nervous centres, or to 
their local action on the lung-nerves, it is difficult to say. I 
believe the latter view the more probable. The sudden cessa* 
tion of respiration, and the marked nervous symptoms that 
follow the injection of these salts into the arteries, are owing 
to the sudden and great pressure that the nervous centres must 
be subject to when the arterial tension is so rapidly increased. 
The smallness of the quantity of these salts, which mixed with 
the blood causes such striking reactions, places them amongst 
our most powerful reagents: in fact, the chloride of palladium, 
in its power in modifying the properties of living matter, sur- 
passes every other inorganic compound with which I have ex* 
perimented. That half a grain of this salt diffused through the 
whole mass of the blood should give rise to such striking effects 
shows the excessive delicacy of living substances as reagents for 
certain inorganic compounds. ^ 

1 In my earlier experiments with these salts, the sndden fall of the pressnre 
in the arteries, and the presence of scarlet blood in the left caYities of the heart, 
led me to oonclade that they paralysed the heart. 



SOME ADDITIONAL VARIATIONS IN THE DISTRI- 
BUTION OF THE NERVES OF THE HUMAN 
BODY. By Fbofessob Turneb. 

I HATE on several occasions recorded examples, which have 
come under my notice in the dissecting room, of variations 
from the usually described arrangement of the nerves of the 
human body {Natural History Review, 1864: this Jowmal, 
November^ 1866 : Proc Roy. Soc, London, 1868). I propose now 
briefly to note some additional variations which I have met 
with. 

Branches of the 5th Cranial Nerve. Left orbit in which the 
frontal nerve gave origin to a long slender infra-trochlear 
branch, which passed below the pulley to be distributed along 
with the infra-trochlear of the nasal. 

Right orbit in which the lachrymal branch of the ophthal- 
mic was not seen, but the temporal branch of the orbital 
branch of the superior maxillary division sent a large branch 
to the lachrymal gland. On the left side of same subject a 
small lachrymal nerve was present, but the temporal branch 
from the superior maxillary, as it lay in relation to the outer 
wall of the orbit, gave an accessory branch to the gland. 

Right side in which gustatory and inferior dental nerves 
formed one cord until opposite dental foramen, when the dental 
nerve entered its canaL An additional case to the one de* 
scribed in Natural History Review, in which the mylo-hyoid 
nerve not only supplied its proper muscles, but gave off a 
branch which pierced the mylo-hyoid muscle to join the gusta- 
tory nerve on the distal side of the submaxillary ganglion. A 
slight modification of this arrangement was seen in another 
subject in which the right mylo^hyoid nerve passed under cover 
of the posterior border of the corresponding muscle before it 
bifurcated, when the muscular branch pierced the mylo-hyoid 
to supply it and the anterior belly of the digastric. Qaillet has 
described some closely similar cases. 

Branch of Vagvs, On left side vagus and hypoglossal were 
intimately connected as low as posterior belly of digastric. 



102 PROFESSOR TURNER. 

Here a large funiculus arose from vagus which gave oflf branches - 
to depressor muscles of larynx, and then, much diminished in 
size, rejoined the trunk of the vagus a short distance above the 
1st rib. In this case the fibres of the descendens noni had 
obviously accompanied the trunk of the vagus for some dis- 
tance, and when this branch was given ofiF some fibres proper to 
the vagus probably accompanied it, which rejoined their trunk 
lower down in the neck. Similar apparent origins of the de- 
scendens noni from vagus have been described by myself and 
other anatomists. 

Posterior Divisions of Spinal Nerves. On right side the 
external branches of the posterior divisions of the 2nd and 3rd 
dorsal nerves pierced the vertebral aponeurosis, gave branches 
to the lower fibres of the rhomboideus major, then pierced that 
muscle to end in the lower part of the trapezius. On left side 
the external branch of the posterior division of the 3rd dorsal 
nerve terminated in the lower fibres of the rhomboideus major. 
On both sides the rhomboidei also received their proper nerves of 
supply from the 5th cervical, and the distribution of the spinal 
accessory to each trapezius was normal. 

Anterior Divisions of the Cervical Spinal Nerves. On left 
side the supra-clavicular cutaneous branch of the cervical plexus 
passed through a canal in the clavicle in its course to the inte- j 

gument in front of the greater pectoral muscle. Bight side of 
another subject in which the 4th and not the 3rd cervical nerve 
gave a branch to the levator anguli scapulae. 

Variations in the course of the phrenic nerve and its lowest 
root of origin are not uncommon. Though this nerve usually 
enters the thorax in front of the internal mammary artery, 
sometimes it goes behind. On the left side of one subject it 
passed in firont not only of that artery, but of the subclavian 
vein (see also Quain's Arteries), In one case the root from the 
6th cervical passed independently down the anterior surface of 
the scalenus anticus to join the phrenic at the root of the neck ; 
whilst in another it ran in front of the right subclavian vein, 
and behind the 1st costal cartilage to join the phrenic close to 
the origin of the arteria innominata. 

The arrangement of the cords and the mode of origin of 
the branches of the Brachial plexus are subject to very frequent 



DISTRIBUTION OP THE NERVES OP THE HUMAN BODY. 103 

> variations, some of which I recorded in my former paper in thd 

Natural History Review. Other variations are as follows : more 
than once I have seen the plexus to possess three, and not two 
cords, in its original construction : an outer formed by the junc- 
tion of subdivisions of the 5th, 6th, and 7th cervical nerves ; a 
posterior by the junction of subdivisions of the 5th, 6th, 7th, 
and 8th, and an inner by the junction of a subdivision of the 
8th cervical with the Ist dorsal. In another case the outer 

'^ cord was formed by junction of subdivisions of 6th and 7th : 

the posterior by junction of subdivisions of 7th and 8th ; the 
inner by a subdivision of 8th cervical joining the 1st dorsal. 
The outer cord gave origin to musculo-cutaneous and outer 
head of median nerves; the posterior to subscapular and 
musculo^spiral ; the inner to ulnar and inner head of median. 
The nerve of Wrisberg, internal cutaneous, and anterior thoracic, 
arose directly from 1st dorsal; the supra-scapular from 5th 
cervical, and the circumflex by the junction of almost equal 
branches of the 5th and 6th cervical nerves. In another case 
the brachial plexus had only two cords in the axilla: a posterior, 
which gave origin to its usual branches; an external, from which 
arose all the branches which usually spring ifrom the outer and 
inner cords. The branches for the inner side of the limb 

^ passed superficial to the arterial trunk. In another subject the 

7th cervical nerve did not join the upper cord of the plexus, but 
divided into three branches, one passed to the inner, another to 
the posterior cord, and the third entered directly into the forma- 
tion of the median nerve. In another subject the outer cord of 
plexus gave origin to both anterior thoracic nerves, to the outer 
head of the median, the musculo-cutaneous, and to a long 
slender branch which crossed obliquely in front of the axillary 
artery to join the ulnar. In another, the anterior thoracic nerve 
which supplied the pectoralis minor, arose by two roots, one 
from inner, the other from outer cord of plexus. In another 
the left ulnar nerve had two roots, one from inner, and an ac- 
cessory funiculus from outer cord, which was traced upwards to 
7th cervical nerve : it crossed in front of the axillary artery and 
joined the ulnar opposite the lower edge of the tendon of sub- 
scapularis. 

Various modifications in the arrangement of the nxedian 



102 PROFESSOR TURNER. 

Here a large funiculus arose from vagus which gave oflf branches 
to depressor muscles of larynx, and then, much diminished in 
size, rejoined the trunk of the vagus a short distance above the 
1st rib. In this case the fibres of the descendens noni had 
obviously accompanied the trunk of the vagus for some dis- 
tance, and when this branch was given off some fibres proper to 
the vagus probably accompanied it, which rejoined their trunk 
lower down in the neck. Similar apparent origins of the de- 
scendens noni from vagus have been described by mjrself and 
other anatomists. 

Posterior Divisions of ^nal Nerves. On right side the 
external branches of the posterior divisions of the 2nd and 3rd 
dorsal nerves pierced the vertebral aponeurosis, gave branches 
to the lower fibres of the rhomboideus major, then pierced that 
muscle to end in the lower part of the trapezius. On left side 
the external branch of the posterior division of the 3rd dorsal 
nerve terminated in the lower fibres of the rhomboideus major. 
On both sides the rhomboidei also received their proper nerves of 
supply from the 5th cervical, and the distribution of the spinal 
accessory to each trapezius was normal 

Anterior Divisions of the Cervical J^nal Nerves. On left 
side the supra-clavicular cutaneous branch of the cervical plexus 
passed through a canal in the clavicle in its course to the inte- 
gument in front of the greater pectoral muscle. Bight side of 
another subject in which the 4th and not the 3rd cervical nerve 
gave a branch to the levator anguli scapulae. 

Variations in the course of the phrenic nerve and its lowest 
root of origin are not uncommon. Though this nerve usually 
enters the thorax in front of the internal mammary artery, 
sometimes it goes behind. On the left side of one subject it 
passed in front not only of that artery, but of the subclavian 
vein (see also Quain's Arteries). In one case the root from the 
5th cervical passed independently down the anterior surface of 
the scalenus anticus to join the phrenic at the root of the neck ; 
whilst in another it ran in front of the right subclavian vein, 
and behind the 1st costal cartilage to join the phrenic close to 
the origin of the arteria innominata. 

The arrangement of the cords and the mode of origin of 
the branches of the Brachial plexus are subject to very frequent 



DISTRIBXrnON OF THE NERVES OP THE HUMAN BODY. lOS 

variations, some of which I recorded in my former paper in thd 
Natural Sistory Beview. Other variations are as follows : more 
than once I have seen the plexus to possess three, and not two 
cords, in its original construction : an outer formed by the junc- 
tion of subdivisions of the 5th, 6th, and 7th cervical nerves ; a 
posterior by the junction of subdivisions of the 6th, 6th, 7th, 
and 8th, and an inner by the junction of a subdivision of the 
8th cervical with the 1st dorsal. In another case the outer 
cord was formed by junction of subdivisions of 6th and 7th : 
the posterior by junction of subdivisions of 7th and 8th ; the 
inner by a subdivision of 8th cervical joining the Ist dorsal. 
The outer cord gave origin to musculo-cutaneous and outer 
head of median nerves; the posterior to subscapular and 
musculo^spiral ; the inner to ulnar and inner head of median. 
The nerve of Wrisberg, internal cutaneous, and anterior thoracic, 
arose directly from Ist dorsal; the supra-scapular from 5th 
cervical, and the circumflex by the junction of almost equal 
branches of the 6th and 6th cervical nerves. In another case 
the brachial plexus had only two cords in the axilla: a posterior, 
which gave origin to its usual branches; an external, from which 
arose all the branches which usually spring from the outer and 
inner cords. The branches for the inner side of the limb 
passed superficial to the arterial trunk. In another subject the 
7th cervical nerve did not join the upper cord of the plexus, but 
divided into three branches, one passed to the inner, another to 
the posterior cord, and the third entered directly into the forma- 
tion of the median nerve. In another subject the outer cord of 
plexus gave origin to both anterior thoracic nerves, to the outer 
head of the median, the musculo-cutaneous, and to a long 
slender branch which crossed obliquely in front of the axillary 
axtery to join the ulnar. In another, the anterior thoracic nerve 
which supplied the pectoralis minor, arose by two roots, one 
from inner, the other from outer cord of plexus. In another 
the left ulnar nerve had two roots, one ftt)m inner, and an ac- 
cessory funiculus from outer cord, which was traced upwards to 
7th cervical nerve : it crossed in front of the axillary artery and 
joined the ulnar opposite the lower edge of the tendon of sub- 
8capularis. 

Various modifications in the arrangement of the median 



104 PROFESSOR TURNER. 

• 

and musculo-cutaneous nerves have been described by Hyrtl, 
Qruber, Qegenbaur, Krause and myself. I may now record a 
case in which the outer head of the median passed behind the 
axillary artery, so that the trunk of this nerve was throughout 
situated on the inner side of the vascular trunk. A case in 
which the musculo-cutaneous pierced both coraco-brachialis and 
short head of biceps, whilst the median passed behind the 
brachial artery. Another in which the inner head of the 
median passed behind the axillary, whilst the trunk of the 
nerve went behind the brachial lower down in the limb. A 
case where the axillary artery gave rise to a large profunda 
branch, between the two vessels the inner head of the 
median passed to join the outer. Lower down the limb the 
nerve went behind the brachial artery. Gruber has noticed 
that in 100 arms the nerve passed 20 times behind the vessel. 
A subject in which a very large musculo-cutaneous nerve 
pierced coraco-brachialis, then gave off a large branch which 
joined the median, but before doing so a branch to rejoin the 
jnusculo-cutaneous lower down in the limb arose from it. After 
giving off the above large branch the musculo-cutaneous a 
second time pierced the coraco-brachialia Another, in which 
the musculo-cutaneous, after piercing the coraco-brachialis, di- 
vided into two branches, one to supply biceps and brachialis 
anticus and to become the external cutaneous nerve of forearm, 
the other to join the median, and also to give rise to a branch 
which joined the external cutaneous 3 inches above the elbow. 
Another, in which the musculo-cutaneous gave off in middle of 
upper arm a fine branch, which descended along with brachial 
artery, and was lost in the deep loose areolar tissue in front of 
the elbow. 

In the Natural History Review I described a case in which 
the teres major derived its nerve from the circimiflex, which 
also supplied the deltoid and teres minor. I have since seen a 
second case of the same kind : but in it 3 subscapular nerves 
were also present, two for the subscapularis, one for the latissi- 
mus dorsi. 

Gruber has recently recorded cases of communication be- 
tween the median and ulnar nerves in the forearm. In January, 
1868, 1 saw a branch arise from the median in the middle of 



DISTHIBUTIOK OF THE NERVES OF THE HUMAN BODY. 105 

the left forearm, and pass obliquely downwards and inwards 
between the flexor muscles of the fingersi to join the ulnar. 
In the same arm the posterior interosseous nerve did not form 
the so-called ganglion behind the carpus, but passed down the 
back of the hand to the clefb between the middle and index, 
divided there into two branches, which supplied the web, and 
ran down the adjacent sides of those digits. Another subject, 
in which the dorsal branch of the ulnar nerve ended in the skin 
of the inner margin of the hand, and the radial formed the only 
dorsal nerve for the digits. A case in which the median sup* 
plied the 1st, 2nd, and 3rd lumbricales^ the ulnar only the 4th. 

Anterior Divisions of Lumbar Nerves. A subject in which 
obturator nerve arose from 2nd, 3rd, and 4th lumbar nerves. 
Another in which the Bcce^sorj obturator descended behind the 
pectineus, and in front of the horizontal ramus of the pubis 
and the adductor longus, to join the obturator and internal 
cutaneous nerves at the inner side of the thigh below the in- 
sertion of the adductor longus. 

Anterior Divisions of Sacral Nerves. Correlated with du- 
plicity of the pyriformis muscle the great sciatic nerve is also 
subdivided into two parts, one emerges below the pyriformis, 
the other between its two subdivisions. In a case of this kind 
I saw the small sciatic also subdivided into two parts, one 
passed below the muscle, the other between its two halves, to 
join the lower root, after it had given origin to the long puden* 
dal and cutaneous branches to the inner side of the thigh. 

Great variability is displayed in the distribution of the digi- 
tal nerves to the dorsum of the foot. In my former paper I 
stated that it largely depended on differences in the number of 
toes supplied respectively by the anterior tibial and external 
cutaneous, but further observation has shown me that the ex- 
ternal saphenous participates in ^hese variations, as I have 
several times seen this nerve supply not only the skin of the 
outer side of the 5th toe, but its inner side, and even send 
digital branches to the 4th and 3rd toes. 

I have not noted here the many variations in relative size 
exhibited by the different nerves which pass to given cutaneous 
areas, as all my observations bear out the view I had expressed 
on this subject in the Natural History Eeview, But I may 



106 KIOF. TURNER. DISTRIBUTION OF THE NERVES, &C. 

state further, in connection with a remark made by Mr Darwin 
in ''the Variations of Animala and Plants under Domesti' 
ctxtton,'* II. 300, that I believe the diflference in size of the 
nerves passing to a given cutaneous area, to be due to original 
variation, and not to increased use or action. Further, I may 
state that variations in the arrangement of the cutaneous nerves, 
which pass to the dorsum of the hand and foot, are more fre- 
quent than in those which supply their opposite aspects; for 
the palm and sole, which are more especially employed as 
tactile surfaces, seem to need more precise and constant con- 
nection with definite regions in the cerebro-spinal nervous cen- 
tres than is necessary for the dorsal aspects of the terminal 
parts of the limbs. 



ON SOME POINTS IN THE ANATOMY OF A GREAT 
FIN-WHALE {Balamoptera Muscuhs). By John 
Struthebs, M.D., Professor of Anatomy in the Univer^ 
siiy of Aberdeen. (Plate VII.) 

This Whale was found dead in the North Sea off Aberdeen, and 
was towed into Peterhead Bay on 27th June last. I have to 
thank my friend Dr Jamieson of Peterhead for his kindness and 
assistance there. At low tide the carcase lay among the rocks 
on its right side with the belly partly turned up, the head 
almost quite on its back. The plaited breast at once showed it 
to be one of the Borquals. The length, measured along the 
back was 68 feet, along the side 64 feet. I take the latter as 
the true length. From the remaining colour, the size of the 
whalebone and its white colour on the bristly surface, the nar- 
row form of the body and head, the relatively small pectoral fin, 
the well marked dorsal fin and its position, the thinness of the 
blubber, and from other characters which will be noticed in the 
course of the paper, it appears that this Whale presents most 
nearly the characters given as distinctive of the Razorback, Bar 
kenoptera muecvlue {Pkyealue antiquorum of Gray, Pterobalcena 
communis of van Beneden). In some of its osteological charac- 
ters, however, it will be seen to present variations from these. 
It was a male, and the state of the vertebrsB shows that it was 
passing from the adolescent to the adult state. 

Bony Rudiment of Hind Limb. — ^The discovery by Prof. 
Flower of a nodule of cartilage attached to the pelvic bone in 
S. musculuSf which he regards^ as the representative of the hind 
limb in this Whale, gave an additional interest to the inquiry 
as to the existence of this rudiment in Fin-whales. After mak- 
ing the external measurements, I dissected out the pelvic bone 
on the exposed side, keeping well clear of it on the outer side. 
In doing so I was in no small degree gratified to feel in my 
hand a separate bone moving on the pelvic bone. At first it 
felt as if in a capsule, but on dissection this proved to be only 
loose fatty tissue. The form and situation of the appendicular 

* Proe. Zoo. See, ofLondoriy Not. 1S66, p. 704. 



108 PROFESSOR STRUTHERS. 

bone, and its ligamentous connection, are represented in Fig. 3. 
In form it is a flattened ovoid, length 2 inches, breadth 1^, 
ihickness |. Of the length, the inner three-fourths are bony, 
the outer fourth, as represented in the sketch, is cartilaginous, 
and from this a fibrous continuation proceeded outwards among 
the soft parts. The bone lay in the hollow between the iliac 
and pubic processes of the pelvic bone, about two inches from 
the former and close to, or even a little overlapping the latter. 
At the proximal end it was attached by a strong flat triangular 
ligament which passed back in two chief bands, the external to 
the iliac process, checking inward motion of the bone across the 
pubes, the internal reaching obliquely backwards in part upon 
the ischial process, checking outward motion. Being joined by 
a thinner part, both together appear as one large triangular liga- 
ment attaching the appendicular bone to the under surface of 
the pelvic bone near the meeting of its three processes, and, 
therefore, in a position corresponding to the acetabulum. There 
was no ligament connecting it to the pubic process or other out- 
ward connection save the fibrous prolongation already noticed, 
the distal connections and nature of which it will be interesting 
to ascertain when an opportunity occurs of following it out 
among the more distant soft parts. 

This observation added to that of Mr Flower may be con- 
sidered as determining the presence of the rudimentary hind 
limb in B, musculus. In Mr Flower's specimen, also a male and 
three feet longer than mine, the nodule was cartilaginous, while 
in mine it was mostly bony, but variation is to be expected in 
parts so rudimentary. When the Cetacean hind limb was first 
discovered by Reinhardt in a new-bom Greenland Right-whale, 
the femur was partly ossified while the pelvic bone and tibia 
were still cartilaginous, but in the full-grown Whale of the same 
species Eschricht and Reinhardt found an ossified femur 8 inches 
long and an ossified tibia 4 inches long, the femur with synovial 
articulation to the pelvic bone at one end, and similarly articu- 
lated to the tibia at the other*. Some might be disposed to 
regard these bones as representing the distal rather than the 
proximal end of the rudimentary limb, but the nomenclature 

^ Memoir on the Greenland Right-whale (Balana Mysticetut), Copenhagen, 
1S61. Ray Society's tnnslation. 



ANATOMY OF A GREAT FIN-WHALE. 109 

of Eschricht and Bernhardt appears to be the more correct 
as well as simple. Abandoning the first view of Eschricht that 
these bones are analogous rather to the marsupial bones of 
Marsupial animals, they interpreted the two subsidiary bones 
in the Greenland Right-whale as rudimentary hind legs hidden 
in the flesh, and as representing the femur and tibia. Trans- 
ferring this interpretation to the single subsidiary bone which 
Eschricht found in three foetuses and one fuU-grown specimen 
of the Greenland Humpback, they consider it to represent the 
femur in that whale. In like manner the cartilage or bone in 
the Razorback may also be regarded as representing the femur. 
The Pelvic Bone — The form is seen in the sketch, Fig. 3. 
The three processes may be termed ischial, pubic, and iliac, fol- 
lowing the interpretation of Eschricht and Beinhardt, although 
they are ossified firom one centre. Homologous parts may in 
one species be developed as outgrowths, in another indepen- 
dently, and may even vary in this respect in the same species, 
notably so in the case of Cetacean transverse processes, as prepa- 
rations in my collection show. The homology of the three pro- 
cesses of the bone is at least relationally correct and the nomen- 
clature is convenient It is a gently curved bone, over 20 inches 
in direct length (23 with the cartilaginous ends), concavity 
inwards, depth of curve 2 inches, (a) The Ischial process, pos- 
terior, 9 inches in length, is the thickest, and prismatic except 
towards the iliac expansion. By its flattened inner surface it 
attached the cms penis for a length of 6 inches, as indicated in 
the sketch. Its upper surface presents a long rough depression 
at and behind the middle. The end is continued by cartilage 
for an inch. (5) The Pvbic process, anterior and internal, 12 
inches in length, is at first flattened like the iliac part, then 
prismatic along its middle third, and in its distal third flattened 
in the opposite directionv the surfaces inwards and outwards. It 
expands a little towards the end which is continued by cartilage 
for an inch, (c) The Iliac process, external, short broad and 
triangular, rises to a blimt summit, which projects 2J inches 
"beyond the line of the outer margin of the bone, and is 4 inchea 
£rom the opposite edge of the bone. Breadth at the base 5 
inches, close to the summit IJ inch. The whole bone is much 
flattened here, thickness | inch, upper surfax» convex, under 



110 PBOFESSOR STRUTHERS. 

surface concave. The ridge across the back part of the iliac 
process on the under surface attaches the ligament of the femur. 
Behind this there are rough depressions. 

Near the fore part of the base of the iliac process there is a 
foramen piercing through the bone, large enough to admit a 
goose-quill, directed obliquely from the under surface outwards 
and backwards. In the sketch I have carried the ligament and 
rudimentary bone a little outwards in order to show the fora- 
men. It was occupied by fat. That this space may have no 
morphological significance may the more readily be supposed 
when the mode of development of the bone is considered. But 
if in comparing the Cetacean innominate bone with that of the 
quadruped or of man, we consider the ischial and pubic perineal 
rami to be here greatly elongated, forming the sides of the Ceta- 
cean perineal girdle, and the iliac bone (which* in its developed 
state is related to quadrupedal or bipedal support) to be here 
stunted, then this foramen occupies the position of the obturator 
foramen in the developed pelvis. 

Muscles of the Finqers. Flexors and Extensors. 
In 1865 Prof. Flower observed the presence of muscles in the 
forearm of a large Fin-whale {B. muaciUtLa), and directed atten- 
tion^ to the importance of having them more fully investigated. 
These muscles have since been examined in the lesser Fin-whale 
{B, rostrata) on two occasions, by Drs Carte and Macalister of 
Dublin* and by Mr J. B. Perrin'. The arrangement of the 
muscles, as found by these authors, was in several respects dif- 
ferent in the two individuals of the same species. I embraced 
the opportunity of observing these muscles, and as they are 
very interesting structures^ and the large scale of the specimen 
enabled the various points in their anatomy to be made out 
distinctly, I have thought it worth whUe to describe them fully. 
The arrangement found, besides, differs in several respects from 
that found by the above-mentioned authors. Their arrange- 
ment is represented in Figs. 1 and 2. I sketched them carefully 
from the dissection, and the artist has reduced them to ^^ on 
the stone. 

1 P.Z.S, 1866, p. 706. 

■ Traru, Roy. Soc. of London, 1868. 

• P.Z.3.1S70. 



v 
« 



d 



ANATOHT OF A GREAT FIN-WHALE, 111 

(a) Internal, or palmar, aspect. There are three musdes 
on the internal, or flexor, aspect of the forearm. They corre- 
spond to the following muscles in man — ^flexor carpi ulnaris, 
flexor profundus digitorum, flexor longus pollicis. 

1. Flexor carpi uhiaris. This muscle arose by a thin fan- 
shaped fleshy expansion from the olecranon cartilage and idna 
near it. Tapering, it ended in a rounded tendon about the thick- 
ness of the little finger, which passed straight along the free fatty 
space between the ulna and the upper edge of the paddle, and, 
expanding a little, ended by being inserted into the nearest point 
and some way along the ulnar slope of the pisiform Gartilage. 
Length 2 feet; tendon occupies rather more than half of this 
and sends an aponeurotic expansion on the fusiform and fan- 
shaped portions of the fleshy part. In the Pike-whale this 
muscle was found by Drs Carte and Macalister to be inserted 
into the fourth metacarpal bone, by Mr Perrin, to the lower end 
of the ulna. Here its insertion into and termination in the pisi- 
form cartilage is quite distinct. The Pisiform cartilage is flat- 
tened, 5 inches in length, 3 in breadth, and is movable upon 
the carpus and ulna. The homology of this cartilage to the 
pisiform bone, and of this muscle to the flexor carpi ulnaris of 
man, is striking. This muscle can have little or no action in 
flexing or abducting the carpus, and the motion of the pisiform 
is too limited to account for the muscle. It may give tension 
to the pisiform. cartilage as the part which is placed at and 
maintai^ the gr^tesTconvexity ofVhe thin upi^r edge of the 
paddle ; it may give resistance to a flexible olecranon for the 
action of the triceps, or even partially transmit the tension of 
the triceps to the pisiform ; but it seems as if its most likely 
effect will be, like a string within a curtain, to give resistance 
to the soft part of the paddle which intervenes between the 
upper edge of the paddle and the ulna. 

2. Flexor digitorum uhiaris, answering to the Flexor pro- 
fundus digitorum of man. It is the largest of the muscles. 
Fleshy part, length SO inches ; breadth at first 7, diminishing 
to 2 ; thickness above middle nearly 2, diminishing distally to 1. 
Is a single penniform muscle, length of fibres 3 to 4 in. Origins, 
over total length of 27 in., partly from humerus, ligament of 
elbow, olecranon, along slope of shaft of ulna, interosseous tia- 



112 PROFESSOR STRUTHERS. 

sue, and septum between it and next muscle. Form, at first 
flat and expanded on humerus and at first on ulna, then thick 
and narrow, and, with next muscle, sunk into interosseous hol- 
low, so that they do not project beyond level of bones. Tendon 
runs up along ulnar margin and sends a stong aponeurotic ex- 
pansion over muscle, but first 3 or 4 inches purely fleshy. 
Where flesh ceases, tendon is 1^ inch broad and about half as 
thick; gradually expands along distal fifth of forearm to a 
breadth of 3 inches, where exchange with tendon of next mus- 
cle occurs, and finally to breadth of 6 inches, opposite first 
carpal row, just before it divides into four tendons. 

3. Flexor digitorum radialis. This muscle corresponds to 
the Flexor longus pollicis of man, in quadrupeds and quadru- 
mana to the external deep flexor which is in various degrees 
united with the internal deep flexor. Is about J bulk of last 
muscle. Fleshy part, length 21 inches ; breadth at first 2, dimin- 
ishing distally ; thickness over 1 inch. Origins, over total length 
of 18 inches, along proximal J of radius beginning 4 inches 
after elbow, fi-om interosseous tissue, and strongly from fibrous 
septum between it and last muscle. Origin covers at first about 
^ of breadth of radius, and narrows as it passes along the inter- 
osseous hollow. Is a bipenniform muscle, the fibres 2 to 3 
inches in length. Tendon sends aponeurosis over distal f of 
fleshy belly, and is concealed as a partition in proximal third. 
Fleshy fibres of this and last muscle cease opposite same point. 
Tendon 1 inch broad, and \ thick. 

The tendons of the two flexors exchanged fibres in a manner 
which may be compared to the exchange, when it is complete, 
between the corresponding muscles in the sole of the human foot. 
The slip from the ulnar flexor formed more than ^ of the tendon 
to the radial digit ; the radial flexor passed mainly to the first 
digit (which in this four-fingered hand answers to the Index), 
sending about J of its bulk as a slip to the ulnar flexor, which 
went on one side entirely to the next digit, in the other limb also 
in part to the tendons of the two ulnar digits. The oval slit seen 
in the sketch existed on one side only. The four tendons sepa- 
rate in the manner shown in the sketch, the two radial 1 J in. 
earlier than the two ulnar. The tendons of the middle digits 
and that of the radial digit have no insertion into the meta* 



I 



ANATOMY OF A 6RSAT FIN-WHALE. 113 

carpal bones, the hollowed surfaces of which are levelled up by 
fibrous tissue, which is smoothly grooved to form the floor of the 
theca. They begin to be inserted into the first joint, or first 
phalanx, and thereafter are continued to the distal phalanges, 
gradually diminishing, and are attached to every phalanx, not 
by one insertion merely but by a close succession of fibrous 
lameUsp. As the tendons pass along the fingers they give off 
oblique slips to the joints, giving the appearance of a diamond-^ 
shaped expansion at each joint. The tendons of the lateral 
digits are twice the size, though not twice the breadth, of those 
of the two middle digits; they pass obliquely, and, before 
reaching the phalanges, have strong lateral attachments to the 
first joint, that of the tdnar digit laying hold also of the distal 
part of the metacarpal bone, thus maintaining their obliquity. 
The lateral digits have thus the strongest tendons, and they are 
so attached as to give them a special hold on the margins of the 
paddle. The middle tendons opposite the metacarpal bones are 
f inch in breadth, and may be compared to thick leather straps 
bevellel off at the edges. Xhe tendons lie in thec88. These 
begin as soon as the tendons, and fit them, with loose connec- 
tive tissue between as far as the expansion, after which the 
separate tendons and their thecsB present smooth synovial sur« 
&ces^ interrupted on the deep aspect where the insertions com- 
mence. The floor of these theosB on the fingers is formed by 
masses of fibrous tissue, levelling up the hollows of the pha- 
langes. These pass firom cartilage to cartilage much more than 
jfrom phalanx to phalanx, and while they partly serve as liga- 
ments binding the thick epiphysial cartilages to the bones they 
serve to fill up the hoUows so far, and are remarkable instances 
of those stratified periosteal thickenings which are seen at 
various parts of the Cetacean skeleton, the whole periosteum of 
which appears to be peculiarly thick. The tendons themselves 
are 'so extensively connected to the phalanges that they may 
be regarded as partly ligaments. 

No trace was seen of any muscle corresponding to the PaU 
maris longuB found by Drs Carte and Macalister, or of the 
Flexor sublimia digitorum foimd by Mr Perrin, in their dissec- 
tions of the lesser Fin-whale. 

{b) Eetemali or dorsal, aspect* The extensor oommunia 

VOL. YL 8 



114 PROFESSOB STRUTHEBS. 

digitorum alone was present. Fleshy part, length 18 in., being 
more than half length of forearm ; breadth, greatest 3^ in., 
diminishing at middle to 2^, near tendon to 1^; thickness 
about 1 incK Origins, beginning 2 inches below elbow, from 
both bones, along narrow part of ulna for 6 in., along radius for 
16 and more broadly; deeply from the interosseous fibro-adipose 
tissue, and superficially, as in man, from a strong aponeurosis 
which binds it down into the interosseous hollow, this aponeurosis 
lurising as high as the elbow-joint. Also from several fibrous 
septa which dip into the muscle and give origin to bipenniform 
bundles. Fibres 2 inches or more in length. Tendon sent 
thick prolongation over distal half of flesh. Tendon, at first 
1^ in. in breadth, passed along distal f of forearm, and opposite 
middle of carpus had expanded to a breadth of 4 inches trans- 
versely. From this expansion the four tendons proceeded in 
the same order as that of the flexors, the tendon to the radial 
digit first Unlike the flexor tendons, the extensor tendons to 
the lateral digits were weaker than those to the two middle 
digits, but they had the same early marginal insertion, main- 
taining their oblique position. All of the extensor tendons 
were attached to the metacaipal bones as well as to the pha- 
langes. As they passed along, diminishing to the distal pha- 
lanx, they broke up at each joint, as in man, into three, the 
middle portion passing straiglit on whUe the lateral expansions 
laid hold of the swollen joint and the phalanx beyond. In addi- 
tion to these tendons and adhering to them, there were strong 
fibrous bands passing along the metacarpal bones and phalanges, 
filling up the hollows ds on the palmar aspect. 

It may seem at first that these digital muscles, larger than 
the hamstrings in a muscular man. and with their red healthy 
fibres indicating activity, must be of considerable importance in 
the limb, but they are ssmll relatively to the si^e and condition 
of the parts on which they act. The whole fleshy mass is' not 
half so bidky as one of the fingers on which they act; and com- 
paring them with the great mass of flesh in the forearm of a 
naan or quadruped, in relation to their digits, we may see how 
feeble they must be as moving powers. Then there is the com- 
parative inflexibility of the joints,, and the farther restriction of 
motion by the uniform covering of akin and blubber in which 



ANATOMY OF A GREAT FIN-WHALE. H5 

the whole is wrapped. It will satisfy any one of this when he 
finds his whole strength exerted in vain to bend one of these 
large paddles, except near the tip and along the soft ulnar 
margin. The length of the fibres composing the muscles shows 
that there is less than an inch of contraction to spread oyer 
a series of joints extending some three feet in length. The. 
total functional result cannot be more than to give some rigidity 
or resistance to the joints, the tendons serving very much as 
ligaments which the muscles will help to tighten, and we see 
the flexors, which will be used in the chief stroke of the paddle, 
stronger than the extensors, especially on the lateral digits. 
These muscles must, therefore, be regarded as rudimentary 
structures, existing through their mammalian affinities and 
maintained by the stimulus of a low function. I have seldom 
if ever contemplated a dissection which impressed me more than 
that of these muscles when first displayed. Within an exter- 
nally unyielding fin, great red muscles and their tendons stretch- 
ing along the white bones, the obvious correspondence to struc- 
tures in the human limb, their vast size, and yet rudimentary^. 
The Ribs, (a) Sixteenth pair of Ribs, ^mong the Fin- 

^ A few dAjB after I had given the above aoconnt of these mnflcleB at the 
meeting of the British Association at Edinburgh in August last, I had, by the 
Idndnees of my pupil Mr John E. Gamer, then at Fraserburgh, the opportunitjf 
of dissecting a Hyperoodon which stranded in Fraserburgh Bay. Although it is 
supposed that no muscles passing from the forearm to the hand exist in any of 
the Delphinoid Cetacea, I may mention here that they were well developed in 
this Bottlenose. It was a male, 20 feet in length. The flexors were arranged 
much as above described in the Bazorbaok, but relatively to the limb they 
were larger, especially the Flexor communis. The Flexor carpi ulnaris passed 
between a long, curved, and flexible olecranon cartilage and a movable pisifoml 
cartilage. On the dorsial aspect there were two common extensors; the radial, 
largest (the extensor communis of man), sent tendons to digits xi, in, and iv; 
the ulnar extensor (the extensor minimi dLgiti of man, the external common 
extensor of the quadruped) sent tendons to digits iv and v. Slight insertions 
into the carpus as they passed seemed to indicate that at least the extensor carpi 
ulnaris was also included. Digit i, the poUex, received no tendon, the extensor 
slip and flexor bundle as if going to the poUex, stop on the first joint of the 
index and partly reach the terminal ligament of the pollex. The digital joints 
reach completely across the cartilage and allow of greater movement &an m the 
Bazorback, in accordance- with the greater development of the muscles. The 
forearm and finger muscles will probably be found to exist in most if not in 
all Whalebone 'Whales, and in others among the Delphinoid Cetaoea besides 
Hyperoodon. Of Narwhal I have a dissection showing their presence morphea 
logically, but histologically they are throughout represented by fibrous tissue, 
and functionally they are ligaments. I inferred that the Flexor oarni ulnaris 
would be the likeliest to be present, and on looking for it in the padole of the 
common Porpoise I find it is present there; the slu>rt fusiform belly attached to 
fuu immovable olecranon, the tendon attached to a movable pisiform cartilage, 
and previouflly oonnected along its distal third to the edge of the ulna. 

8—2 



116 PROFESSOR STRUTHERS. 

whales B, rostrata has 11 pairs of ribs, B. muscvlus 15, B, 8ib- 
baldii 15 or 16. As a developed 16th pair in Sibbaldius appears 
to have been seen only in the Hull skeleton, it is most likely an 
individual variety. As far as observations go B» muscultis shows 
a tendency to diminution rather than increase of ribs, the 15th 
lib being sometimes found much reduced and placed far from 
the spine, but a small 16th rib attached to the first lumbar 
transverse process has also been seen^. The variety which I 
foimd in this Bazorback was the occurrence of a 16th pair of 
ribs suspended in the position in which a reduced 15th pair is 
sometimes found, far from the spine. The right 30 inches in 
length, the left 22, each end prolonged for an inch by cartilaga 
In form, the lower § vary from IJ to 1} inch in breadth, the 
upper i undulating and tapering to a narrow point Neigh- 
bouring part of 15th ribs 2 to 2^ inches in breadth. They 
were suspended opposite the distal part of the 15th ribs ; the 
soft parts connecting the right were already severed, but the 
left projected six inches beyond the end of the 15th rib. The 
intercostal space was 2 to 2| inches in breadth, and was occu* 
pied by a well-marked external intercostal muscle, without 
trace of an internal intercostal As it is apt to be overlooked, a 
guspended additional rib may possibly, be not unfrequent in 
Cetacea'. The 15th ribs are well developed, length 72 inches 
presenting the peculiar sinuous and twisted foim described by 
Dr Murie (P. Z. 8. 1865) in his account of the Rosherville 
skeleton. The longest rib measures 114 inches along the outer 
edge, straight 87. 

(b) Separcxte Gapittdar process of First Mib. The follow- 
ing observation may perhaps help to account for the diflTerent 
appean^nces presented by the head of the first rib in Whales. 
Articulated to the end of the first rib, as seen in Fig. 4, there is 
a, sei)arate beak-like bone or procesig, with intervening layer of 
cartilage. This cartilage is now broken across, but the connect- 

1 See various mention of the Bibs in Whales by Prot Flower, NoU$ on the 
SkeUtont of WhaUt in the pnnoipdl Museums of Holland and BelMum, P. Z. 8. 
1864 ; and in P. Z, 8. 1866. p. 472, and p. 699 ; 1869. p. 604 ; 1870, p. 880. 

' Bmoe the above was wntten, I have noticed in a oommon Porpoise, a male. 
5 feet long, the Idth pair of ribs suspended in exactly the same position as the 
16th in this Bazorbadc. the short 8 inch rib suspended behind the inferior paxt 
of the 8 inch vertebral 12th rib. A ligament passed np from it for 8 inohea 
towards the spine, where it had been divided in separating the ribs. 



ANATOMY OF A GREAT FIN-WHALE. 117 

ing ligaments still remain in the preparation. It fits aocurately 
against the surface of the rib from which it has been detached. 
Length 4 J inches, tapers to a blunt end, inner border convex, 
outer border concave, front surface convex, hinder sur£ace flat. 
At base, breadth If in., thickness 1 ; at middle, breadth 1^, 
thickness ^ inch. Unfortunately the connection of the first 
ribs to the vertebree was severed in my absence and imder 
circumstances which must leave it in doubt whether this pro- 
cess was present on the right side also. The ankylosis of such 
a process would give the rib a well-marked capitular beak, 
while its loss, in forcible disarticulation or in maceration, would 
leave the rib with the form usually represented. The upper 
part of the second rib is represented in Fig. 5, showing its well- 
marked capitular process^ 10 inches long. Length of entire rib, 
along outer edge 93 inches, straight to angle 68, straight 
to head 61, depth of curve 19. Breadth, below angle 4f , average 
of shaft 5|. First rib* Length, straight 51^, along outer border 
64; depth of curve 11| ; breadth between tubercle and angle 
b\ to 6, below tubercle 4f , half-way to sternum 6J, close to 
sternum 6, at lower end 11, including the cartilage 12. 

The Sternum. — ^This sternum is interesting as showing 
variation, and as bearing on the question of the differences be- 
tween that bone in B. musculus and in B, SibboMii. The form 
is shown in Fig. 4, sketched from the preparation, 4 feet 3 inches 
in each direction, after it was carefully cleaned, the natural con- 
nection between the sternum and ribs being left. The sternum 
and first rib articulate at two places, externally (lateral costo- 
stemal articulation), for 5 inches, below the outer part of the 
wing, and internally (terminal costo-stemal articulation) in the 
recess between the wing and posterior process. These articular 
tions are effected by fibrous tissue, and there is a cartilage at 
one side of each, on the sternal side of the lateral, and on the 
costal side of the terminal joint These may be regarded as 
cartilages of growth, but they also give elasticity at the joint 
and attach the ligament. Only the interosseous part of the 
ligament is shown in the sketch, the more superficial and 
stronger part of the great connecting ligament having been dis- 
sected off to show the exact relation and form of the bones and 
the cartilage. The motion was free at both joints, the interoo** 



118 PROFESSOR STRUIHE^S. 

seous part of the ligament being about half an inch in lengthy 
but the two joints assist each other and give a very strong 
union. Between the two joints is a space 6 inches in breadth. 
The sternal end of the first rib expands greatly and presents two 
separate cartilaginous tips^ between which the middle third of 
the end of the rib is thin^ concave and non-cartilaginous; one on 
the anterior angle or process, 3^ to 4 inches broad, belongs to 
the terminal joint, the other at the posterior angle is free. The 
deep recess between the ends of the first ribs is 9 inches broad 
at the middle, 8^ at the hinder end. It is 8|^ to 9 inches in 
depth at the sides, to the sternal joint, subdivided by the xiphoid 
process which projects free into it, and stops short 4 inches from 
the mouth of the notch. 

It is in the length of the posterior process and in the greater 
or less filling up of the antero-lateral notches that these sterna 
vary. Viewed in its connections, the interpretation of the es- 
sentral parts of the sternum is evident. The thickest part 
reaches across as a beam opposite the costal joints, supporting 
the first ribs and completing the visceral arch. The hinder part 
of this beam is at the root of the posterior process and between 
the two terminal joints: the anterior part of it is between the 
lateral joints, a^d forms the posterior ^rtion of the wings. In 
front of this the bone begins to shelve o£f rapidly to a thin 
cervical edge, the anterior development of the median process 
and the wings and the greater or less filling up of the antero- 
lateral notches, depending on muscularity, age, and individual 
variation. At the deepest part of the postero-lateral notch, at 
the meeting of the posterior pedicle with the wing, there is a 
special articular notch, most marked on the pedicle, for the ter- 
minal joint of the rib, and so far the pedicle is essential and ex- 
ists in all the varieties. The pedicle may stop here or project 
variously beyond it in diflFerent species and in different indivi- 
duals, depending on muscularity, age, and variability. 

The sternum in the Razorback is known to vary in its form 
and dimensions*; but in this Razorback it departs so much from 
the form assigned to it as distinctive from that of Sibbaldius as 

* lilljeborg, SynopfU of the Cetaceotu Mammalia of Scandinavia, 1S61-2, 
Ray Sooiety^B translation; and see dimensions of specimens given by varions 
antiioni. 



V" 



ANATOMY OF A GREAT FIN-WHALE. 119 

to have on the ii?hole a greater resemblance to the latter, more 
80 anteriorly, leas so posteriorly at the xiphoid process, which is 
intermediate. This will be seen if Fig. 4 be compared with the 
figures given of the sternum of the Bazorback and of Sibbaldius 
by Prof Flower* and by Prof Turner" of his Longniddry Sib- 
baldius, the latter being firom a very large and nearly adult spe- 
cimen, the length and breadth of the sternum in it and in this 
Sazorback being exactly the same. 

The following are the measurements of the sternum, in 
inches. Breadth, straight, 26^, add over an inch at each end for 
cartilage ; length 17^ ; length of anterior median process, firom 
level of notches, 4; depth of antero-lateral notches 2^; length of 
wing at narrowest part 6^, at broadest 6^; posterior process, 
length 7i, from level of wings 8^, breadth where first ribs meet 
it 4|, two inches from apex 2 ; depth of postero-lateral notches, 
right 4f , left 4^. Thicknesd at mesial line on level with back 
part of wing 2^; back part of wing at middle If, wing in front 
of this rapidly shelving; base of median cervical process If; pos- 
terior process, at base 2^, at middle 2^, before bevelling of point 
If. Foramen, from end of anterior process 7j^, passes through 
the bone very obliquely forwards and upwards, admits goose- 
quill. Depth of concavity along median profile of bone If, but 
processes a little convex, especiaUy posterior; transverse con- 
vexity of bone gives upward retirement of 6 inches opposite 
outer end of wing. On visceral surface, depth of hollow from 
transverse concavity 3^; division of bone, longitudinally, into 
three parts well marked, middle thick and flat, anterior third 
shelving forwards and concave both ways, posterior third, on 
pedicle, a little concave both ways. The general view of the 
pectoral surface suggests a saddle. The prominent longitudinal 
ridge is 1^ inch to right side of middle line, and anterior process 
is correspondingly twisted. Whole of anterior edge of wing 
thin, sharpest at the hollow, becoming rough at the convex part. 
Hinder edge sinuous and rough at gap between the joints. Pos- 
terior process prismatic, lower edge of notch for terminal costal 
joint well marked, process tapers behind this, at first rapidly 
then gradually; last IJ inch rapidly bevelled to blunt point, 
rough, and was coated with cartilage. 

» P. Z. 8, 1864, p. 898. « In this Journal, May 1870, p. 378. 



120 PROFESSOR STRUTHERS. 

Cervical Yertebrje. — ^Among the osteological characteis 
of tlie Bazorback have been given — " Neural arches of the cer- 
vical vertebrae low; spinous processes very slightly developed" — 
and among those of Sibbaldius " Neural arches of the cervical 
vertebrae high, and their spines well developed \*' I notice this 
point as I have the neck of another great Fin- whale, a larger 
and older male, ashore near Wick in 1869, and the comparison 
of the two shows marked difference in the above character. In 
the Peterhead specimen the arches are comparatively high, 
giving a tnangular form to the canal, with well-marked though 
not long spines. In the Wick specimen the arches are so low 
and broad as to give the canal rather a semilunar form, the 
spines are wanting, or are mere roughnesses, on the three mid- 
dle vertebras and very short on the 6th and 7th; the processes 
which project backwards from the laminae, internal to the pos- 
terior articular processes, are much better marked than the 
rudimental spines; and the laminae are, anteriorly, as thin as 
writing paper, in some parts cribriform and flexible, while in 
the Peterhead specimen the laminae are strong. The height of 
the canal in the latter is partly owing to the upper wall of the 
bodies being as yet concave transversely, while in the Wick spe- 
cimen they have become convex, but the lowness of the arch in 
the latter is mainly owing to the pedicles being placed farther 
out on the bodies and to the laminae being low. The thinness 
of the laminae would seem to indicate that disappearance by 
absorption has been going on. Nor does the supposed distinc- 
tion in the transverse processes of the atlas, and in the spine of 
the axis, seem to be reliable, the differences not being greater 
than muscularity, size of animal, and age would account for. 
This will be the more readily understood when we call to mind 
the considerable variation presented by the spine of the axis 
and the transverse process of the atlas in man. 

The rings of the transverse processes are complete from the 
second to the fifth, but the terminal expansions external to the 
apertures are as yet very little developed in the 3rd and 4tb. 
The 6th presents the not unfrequent bony deficiency in the 
inferior transverse process. It occurs here on both sides, at the 
grooved part, for about 1| inch. The gap was filled naturally 

1 Flower, P. Z. 8. 1S64, p. 302. 



ANATOMY OF A GREAT FIN-WHALE. 121 

hj cartilagmoiis tips and thick intervening ligament. This gap 
exists only on the right side in the older Wick specimen. The 
7th vertebra, as usual^ has no inferior transverse process. 

Characters and Measurements of Various Parts. — 
The following notes of characters and measurements will show 
the species to which this Whale belonged, and may be of interest 
to the anatomist for comparison with the characters in other 
specimens. All the numbers refer to inches unless feet are 
mentioned. 



(a) Colour, Putrefiietion having akeady commenced, the colour 
could not be much relied on. It was black on the back, and patches 
of adhering dark cuticle here and there on the sides and in the 
furrows showed that the dark colour had extended well down the 
sides. No traces of dark colour were seen on the mesial ridges or 
furrows, as if the colour had been naturally white there^ Unless the 
carcase is fresh and moist, statements in regard to the colour of 
Whales must be received with caution. The blubber was about 
2 inches thick. 

(b) Tail Fin. Tip to tip 14 feet. Greatest antero-posterior 
measurement of each side 3 feet 8 in. The surfaces and edges were 
so worn and macerated as to present a fibrous appearance, so that the 
tail might readily have been described by a person unacquainted with 
structure, as covered with white hair. With regard to the view that 
one fluke is naturally bent up and the other bent down, on the prin- 
ciple of a screw-propeller, J am led to put the question, for future 
observers, whether this difference on the two sides is not the acci- 
dental result of position. Here the right fluke was concave up, the 
left convex up, and this was evidently owing to the body lying on 
the right side. The carcase usually lies more or less on one side, 
the natural effect of which on a transverse tail is to render that fluke 
concave up, the other convex up, and lying some time in this position 
will perpetuate it in drying. Mistakes are apt to be made in report- 
ing as to right and left. 

(c) Doraal Fin. Well marked. Falcate. Height vertically 15, 
length at base 24 to 26. Distance from end (median notch) of tail 
15 feet 8 in. Distance (of middle) behind anus 28. Near the base 
of its concave posterior margin there was a deep semilunar notch, 
two inches in diameter. Though now perfectly regular and smooth- 
edged, it may have been the result of a bite or shot The i-emains of 
a rocket, which the whaJe-flshers believed to have been the cause of 
death, was found in the right pelvic region. 

(d) Caudal region of Trwnk. From anus to end of tail 18 
feet^ to anterior edge of tail, being caudal part of trunk, 15 feet. 
Presents strongly the ^ razor-back " character, suggesting almost a 
double-edged knife. This much more strongly marked along the 
posterior than on the anterior half. Dorsal ridge farther from lateral 



122 PBOFESSOR STRUTHEB& 

line than ventral ridge and more convex. Semi-girths of this region,^ 
at anus 6 feet 4 in. ; \ way back, 5 feet 5 in. ; halfway, 5 feet; j way 
back, 4 feet 4 in. ; halfway between last and tail, 3 feet 9 in.; just in 
front of tail, 2 feet 10 in. Median lidge runs half way back on tail 
fin. 

(e) These semi-girths may be relied on; those taken over abdo- 
men or thorax are liable to be affected by distention. Approximative 
semi-girth opposite hinder end of pectoral fin was 13 feet. Semi-girth 
two feet behind eye, 13 feet 10 in., just behind eye, 13 feet 7 in. The 
jplaitifigB were mostly fully opened out, the fan*owB being now as 
broad as the ridges. They commenced at side of lower jaw 4 feet in 
front of eye; extended back 32^ feet on the side, 2 feet farther on 
the belly, reaching on the belly to 6 feet from penis, which was pro- 
truded at 4 feet in front of anu& 

(f) ffecuL Eye, Distance from beak of upper jaw to anterior 
canthus, along the lip 13 feet, straight 12 feet 9 in. Canthus to 
canthus 4^. Ear-hole^ — Behind posterior canthus 30 inches, in front 
of where upper edge of paddle leaves trunk 66. Admits with diffi- 
culty point of little finger, cuticle now off ; 4 to 5 inches in, not 
larger than crow-quilP. Projection of lower jaw beyond upper 
about 2 feet, but exact distance uncertain as lower jaw was thrown to 
one side. From beak of upper jaw, along whalebone, to deep part of 
angle of mouth 12 feet 4 in. Superficial part of angle is 9 inches 
farther back. Whalebone, — On outer view black, hairy tips showed 
a white fringe; whole of inner, or buccal, surface, a white hairy 
matting, not cream-coloured as in Pike Whale, but quite white. 
Length of longest plates on outer side, measured straight, 22 inches, 
at symphysis 6. The two sides continuous at symphysis. Breadth 
of the grooved surface of the whalebone matrix, after the plates were 
removed, mea^iured straight, the surface being concave across as well 
as longitudinally — at 6 inches from front, 2 inches; one foot back, 5; 
two feet back, 8; three feet, 10^; four feet, 12; from six to eight 
feet back, 13; ten feet back, 12; ten and a half feet back, 11; twelve 
feet back, 9. Breadth of rim of jaw outside whalebone, at side 6 
inches, at symphysis the same. FalcU&. — Like a raised beam, narrow 
and smooth, convex transversely for 17 inches back; then a median 
groove broadening backwards to six feet back, behind which palate 
projects more and more to a rounded margin. Breadth of free 
rounded surface, at front 6 in. ; after six feet back, narrows to 5, 
then to 4. Total length, from 6 in. behind beak, 12 ft. 4 in. Height 
to which palate projects above level of whalebone matrix — at three 
feet back, 3 in.; at five feet, 6; at seven feet« 9; at nine feet, 13 td 
14; at ten feet, 15; at eleven feet» 14; at twelve feet, 6. Total 
length of akullf from beak of upper jaw, 15^ feet. Greatest breadth of 
skull, just behind temporo-mandibular cushion, 6 ft 3 in. The foUow- 

^ The catiole being o£f I could not ascertain whether there was present the 
fine white streak which I obserred in a 14A feet long Pike Whale, running 
forwards from over the shoulder to the ear-nole, which was very small and 
grooved posteriorly, so that in swimming forwards the water will not enter. 



AHATOMT OF A GREAT BTO- WHALE. 123 

ing breadths of the beak had to be taken from below, across roof of 
mouth) and therefore in a double inclined plane. The rise of the 
palate backwards will be kept in mind as one cause of the increase 
in the measurements backwards. At one foot baok^ 17 inches; at 
two feet, 23; at three feet, 28; at four feet^ 34; at five feet, 38; at 
six feet, 42; at seven feet, 43; at eight feet, 45; at nine feei^ 48; at 
ten feet, 50. 

TemporcHncmdibuhr cushion. A fibro-adipose mass^ now mea- 
sures, in length 30 inches, in breadth 24, in height 15 to 18; lower 
jaw haTtng been separated, the mass may have been larger. Front 
half light yellow colour, more fattj, cuts like firm blubber; hinder 
half white and yellow colours mixed, more dense, the fibrous element 
predominating; transition from front to back gradual. Lower Jaw. 
— Length, straight 14 ft. 4 in.; along curve on outer side 15^ 
feet. Depth of curve at middle, where greatest, 2 feet. Body, height 
near symphysis 9 in., at middle 12; greatest thickness 7. Lower 
border sharp all along; upper border, or surface, 6 to 7 inches 
broad, its inner edge rising up sharp along the three feet next 
coronoid process. Height of bone to tip of coronoid 21, at one foot 
in front of coronoid 15. Height of coronoid process proper, behind, 
from level of front of sigmoid notch, 9 ; in front, from level of body, 
4A; intermediate, 7. Across its oblique base, 9; across middle, 6. 
Thickness about 2|. Highest part of upper edge is in front, from 
which it slopes down and back. Upper half bent outwards. Jaw 
between condyle and coronoid concave externally, convex internally. 
Length of sigmoid notch to tip of coronoid 30, to back of coronoid 24 ; 
greatest depth of notch 8. Dental foramen, from condyle 17, from 
coronoid 1 1. Contents : mouths of at least 24 vessels seen ; one 
artery ^ inch in diameter, two less, and other four or five larger than 
goose-quill; the others, size of small goose-quill. Veins, the thin-coated 
vessels, much smaller than the arteries. A large nerve, as thick as 
thumb, funiculi in a sheath, placed above vessels. Temporo-mandi^ 
bular cushion adhers to condyle. Uniting tissue at symphysis, about 
2 inches thick between, thicker in front superficially. 

(g) Pectoral Limb, Distance from beak of upper jaw to shoulder 
19 feet^ to axilla 21^ feet. The form is represented in the sketches 
Figs. 1 and 2. The small size of the paddle was evident on the first 
view of the carcase and characteristic; length along lower border 
8 feet, along upper border 5^ feet. Exact length after removal, 
from head of humerus, 7 feet 8 in. ; greatest breadth, at the pisiform 
cartilage, 19 in. On dissection the bones and joints presented the 
following measurements, in inches. ScapiUa, height 28; breadth 51, 
increased by posterior cartilage to 62 ; a strip along base connects 
the two cartilages; anterior cartilage adds 4 to height near anterior 
part of base. Coracoid, length 5^, increased by cartilage to 8, 
breadth 3|. Acromion, length 10, increased by cartilage to 11^, 
breadth, 4^. Glenoid ligament much deeper at hinder end of cavity, 
and capsular ligament continuous with it. Humenu, Both epiphyses 
ankylosed, length 21, including cartilage, which is -^ thick on the 
head. Same over all the bones at elbow, but thicker at the edges and 



124 PBOFKSSOB STRUTHEBS. 

ridges. LengUi of Eadius 33 ; of Ulna, from elbow, 30^, from end of 
bony olecranon 37^. Olecranon cartilage^ projection along middle 10, 
antero-posteriorly 10^; thickness near bone If, at middle 1, shelving 
to edges, distal half flexible towards surfaces but not longitadinallj. 
Especially the thicker pai*ts marked with numerous pits. 

Carpite, At middle the soft mass is 7 to 8 in length. Included 
in this are (a) Epiphysial cartilages of radius and ulna 1 to 1^, about 
half of these conceal the bony epiphyses at depth of { inch, half pro- 
jects beyond, (b) Epiphysial cartilages of metacarpals, those of the 
middle digits |. (c) The carpus proper, five cartilages (beside pisi- 
form) with their central ossifications, 3 in first row, 2 in second. On 
palmar aspect the 3 bones of first row nearly of same sise, those of 
second row considerably smaller. Od dorsum, first row nearly as on 
palmar, scaphoid lesa^ cuneiform more; in second row magnum more, 
unciform much less. The true anatomy of the carpus is seen on ex- 
amining the cartilages of which these '' bones " are the more or less 
ossified centres, leaving, in this specimen, variously half an inch or 
more around of cartilage unossified. When the perichondrium is 
removed the outlines of the cartilages are recognised, united along 
their adapted articular surfaces by narrow tracts of ligament First 
row 4 as in man. Scaphoid and semilunar articulate with radius, 
cuneiform with ulna. Pisiform articulates distally with cuneiform, 
at middle with ulnar epiphysis, and for an inch by loose ligament 
with bony ulna; length 5, free edge 3^, breadth 3; thickness near 
articulated edge ^, at middle f , shelving to edges ; flexible on sur- 
faces ; motion at articulated edge, but very little in direction of flexor 
carpi ulnaris. Second row, must be regarded as homologous with the 
magnum and unciform. Magnum senal with digit III, articulates 
also with part of II and part of lY, as in man. Unciform, serial 
with lY and articulates also with part of Y. Two-thirds of Y rest 
on cuneiform, and half of II on Scaphoid^ The lines of articulation 
of the carpal cartilages are shown in Figs. 1 and 2 as fiir as the ten- 
dons and the great reduction permit. The similarity of the articu- 
lations of these cartilages to those of the corresponding bones in 
man, and other mammals, is evident. 

Digital Bones, Number in each of the four digits, including 
metacarpal, II and Y, 4 each; III 7, lY 6. As the lengths of the 
metacarpals and phalanges are given as distinctive between B. 
Musculus and Sibbaldius, I subjoin the length of each, in inches, 
taken at the middle, in their order. II 5, 5|, 5, 3^, terminal car- 
tilage 1^; III 6^, 6^, 5, 34, 2^, 1^, |, terminal cartilage 1; lY 5^ 
^41 ^h ^i' ^i> ^' terminal cartilage 1 ; Y 4}, 4f, 3}, 1^, terminal 
cartilage 1. Total length of each finger before the cartilages had 
shrunk— II 24; ; III 33^; lY 31; Y 19. The cartilages also were 
measured, but it may suffice to mention generally, that the proportion 
of cartilage (iucluding the terminal cartUages) to bone in the fingers 

^ In the Wick speoimen there is a small separate bone in the sitaation of the 
trapezoid, serial with 11, not showing on the snrfaoe. Also, the msgrnm and 
unciform have uiited, on the two su^aces but not deeply, into one bone* 



ANATOMY OF A GREAT FIN-WHALE. 125 

I 

^ is about one to three, cartilage fomiing about one-fourth of the whole 

digit The swollen "joint" between the successive phalanges is a 
mass of cartilage adherent to the bones, and continuous from bone to 
bone except partly across the middle, where, for the middle third or 
less, there is a narrow joint, seen on both aspects^ covered in by 
membrane. This joint disappears at the smaller interphalangeal 
cartilages. The mode in which the nodes and hollows of the two 
radial digits are adapted by alternation is shown in the sketches. 

I r^ret that circumstances prevented a more complete ex- 
amination of the anatomy of this Whale from being xnade. The 
difficulties attending the dissection of so large a carcase, the 
more so when it is already putrid, are well known to those who 
have tried. The weather while I was working on the beach 
was not favourable, and most of the remaining work had to be 
done in the month of July, when I was occupied with the duties 
of the summer session. I must not omit to mention that in 
making out these dissections I derived valuable help from my 
experienced and accomplished Assistant, Dr James Bodger. 



I 



ON THE RELATION OF THE TEMPERATURE OF 
THE AIR TO THAT OF THE BODY. By A. H. 
Garbod, St John's College, Cambridge. 

The nature of the evidence affecting theories in biological 
science is generally so far from direct, that it is only by the 
systematic working out of many of the necessary deductions 
that any idea can be formed as to their value. The experi- 
ments detailed in this communication were suggested by the 
theory to be referred to immediately, and their close agree- 
ment with its requirements tends strongly to substantiate its 
accuracy. 

In a paper published elsewhere* I have detailed several 
observations which tend to shew that many of the minor 
fluctuations in the temperature of the human body result 
from alterations in the amount of blood exposed at its surface 
to the influence of external absorbing and conducting media. 
Others have repeated these experiments', and obtained the 
same results. 

For example; on stripping the healthy body in an air of 
about 60'F., a rise of the internal temperature (judged by 
that of the floor of the mouth) commences immediately, and 
in about half an hour amounts to a« much as three-fourths 
of a degree. According to the above-mentioned theory this 
phenomenon is explained thus; the contact of the cold air 
against the surface of the skin, previously maintained at a 
much higher temperature by the clothes covering it, produces 
80 considerable a contraction of the cutaneous muscular vessels, 
and the blood is driven so far inwards, that the conducting 
power of the thus modified skin is rendered considerably less 
than that of the clothes and blood-filled skin combined^ in 
the previous condition; consequently the body temperature 
rises until a higher equilibrium is attained. 

Such being the case, and the contraction of the cutaneous 
vessels being evidently caused by the cold, it ifl more than 
probable that the amount of thi^ contraction should depend 
on the degree of cold applied, that is, on the temperature of 
the external air in which the observation is being conducted; 

1 Proe. n. 8, No. 112. 1869, p. 419, et aeq. 

" J. F. Goodhart, Ouy's Hotpital ReporU, 1869. 



KELATIVE TEMPERATURE OF THE AIR AND BODY. 127 



and the extent of this action would manifest itself by its 
effect on the body temperature, less cutaneous contraction 
causing less diminished conduction and consequently less rise 
of temperature on stripping. 

Similar reasoning would lead us to anticipate a temperature 
of air sufficiently high to produce exactly as much cutaneous 
contraction as will make up for the loss of the clothing, and 
consequently no change in the body temperature on stripping. 

The correctness of these deductions may be judged from 
the following observations, which were all made under similar 
conditions, on myself, while standing. 

I. n. 



Time. 


Temperat. \ 


remp. Air. 


11.16 


98.95 •) 


11.20 






11.25 


98.975 




11.80 


98.975 




11.35 






11.40 


99.3 


. 47»F. 


11.45 


99.35 




11.50 


99.575 




11.55 


99.625 




12 maHT 


99.7 




12.5 


99.675 . 





Stripped at 11.30. Bi8eO<>.7F. 

ni. 



Time. 


Temperat. ' 


remp. Air. 


10.45 


98.1 




10.60 


98. 




10.65 


97.925 




llP.X. 

11.5 


«fvl. 

98.19 


■ 69«P. 


11.10 


98.85 




11.15 


98.4 




11.20 


98.425 





Time. 


Temperat. Temp. Air. 


11.16 


98.8 ^ 


11.20 






11.25 


98.8 




11.80 


98.8 




11.35 
11.40 


99. 


► 62»F. 


11.46 






11.50 


99.85 




11.65 






12 NIOET 


99.875 . 





Stripped at 11.80. Rise 00.575 F. 

IV. 



Time. 


Temperat. 


Temp. Air. 


11 P.M. 


^ 




11.6 


99. 


• 


11.10 


99. 




11.15 


99. 




11.20 


99. 


. 670 F. 


11.26 


99.06 




11.80 


99.19 




11.86 


99.21 




11.40 


99.19 





Stripped ai 10.65. Bi8e00.6F. 



Stripped at 11.15. Bi0eOO.2F« 



128 



MR QARROD. 



VI. 



Time. 


Temperat. 


Temp. Air. 


10.16 


99.15 




10.20 


99.15 




10.25 


99.16 




10.80 


99.125 




10.86 
10.40 


99.025 
99.125 


► 71«F. 


10.45 


99.176 




10.60 


99.176 




10.65 


99.176 




11 P.M. 


99.175 





Stripped at 10.80. Bise 00.05 F. 
VIL 



Time. 


Temperat. 


Temp. Air. 


10.80 


99.21 




10.86 


99.205 




10.40 


99.2 




10.46 


99.8 




10.50 


99.8 




10.66 


99.16 


. 72«F. 


llP.Ji. 


99.86 




11.6 


99.8 




11.10 


99.825 




11.16 


99.4 




11.20 


99.426 J 





Stripped at 10.60. BiBeO«.125F. 



Time. 


Temperat. 


Temp. Air. 


10.16 


98.8 




10.20 


98.876 




10.25 


98.95 




10.80 


98.95 


. 71«F. 


10.86 


99. 




10.40 


99. 




10.45 


99. 





Stripped at 10.26. ^ Biae (fiM F. 



ym. 



Time. 


Temperat Temp. Air. 


10.16 


98.8 -^ 


10.20 


98.825 




10.25 






10.80 


98.9 




10.85 
10.40 


99. 


' 78«P. 


10.45 


99.1 




10.60 


99.1 




10.65 


99.1 




11 P.M. 


99.1 J 





Stripped at 10.86. BiseO^^.lF. 



It is readily seen that the hotter the air, the less is the 
stripping rise, and that when an external temperature of 
70^ F. is reached^ there is no rise at all. Several of the 
higher temperature observations are here given and but few 
of the lower, because in the paper above referred to {Proc jR. 
S. No. 116, 1869) there are six or seven of the latter recorded 
with the temperature noted in all* 

From these facts it may be clearly seen that in air bdow 
ih4 temperatttre of 70^ F., ^ stripping rise varies inverseljf aa 



RELATIVE TEMPERATURE OF THE AIR AND BODY. 129 

i ihe temperature. Experiments as low as to 45^ F. have been 

made, but no limit has been reached in that direction yet. 
Above 70* F. of the air, the results are modified by the sweat- 
ing that always accompanies so high a temperature if the 
body is clothed, and quite a different class of phenomena 
appear, which have not been much studied. 

In V. and VII. there was a slight fall of temperature at 
the moment of stripping, this was probably connected with the 
perceptible moisture on the surface which evaporates almost 
immediately the clothes are removed. 

With regarcl to the nature of the clothing removed. It 
always had considerable non-conducting power, being com- 
posed in all cases of at least two layers of woollen material; 
though^ as the observations in the warmer air were made 
in summer, and those in the colder, during corresponding 
seasons; the dress worn varied with the time of year, being 
thinner in the former and thicker in the latter. On the 
whole the amoimt of clothing worn does not seem to affect 
the results as long as there is sufficient to keep the body 
warm under ordinary circumstances; and in the English 
climate to do this, one woollen covering seems always es- 
sential. 

Some results obtained by Dr V. Weyrich* with regard 
to the hygrometric condition of the skin at different tempe- 
ratxires of the atmosphere, obtained by means of an hygro- 
meter speciaUy adapted for the purpose, bear so fully on the 
subject under consideration that they will be here given. 

Ist, When the body is clothed, the amount of moisture 
excreted by the skin, does not vary appreciably when the 
observations are conducted in an air below 70" F. 

2nd, When above 70* F. the amount of moisture excreted 
by the skin rapidly increases with a rise in the temperature 
of the atmosphere. 

It is thus seen that by means of an entirely different 
method of observation, Weyrich finds that in an air of 70" F. 
sweating commences ; and by a combination of his results with 

^ J>U Unmerkliche Wcuserverdunstung der Menschliclien Haul, Leipzig, 
1862. Abstract in BriU and For. Med, Ch. Rev, Oct. 1868. 

VOL. VI. 9 



130 MR GAlRROD. RELATIVE TEMPERATURE OF THE AIR, &C 

those arrived at from the facts given above, the following con- 
clusions may be drawn : — 

With regard to the human body, when covered with badly 
conducting clothing, 70* F. is a critical temperature of the 
atmosphere. The removal of the clothing at that tempera- 
ture produces sufficient contraction of the cutaneous muscular 
arteries to counteract the cooling effects of its loss, and con- 
sequently the internal temperature does not change; whilst 
on stripping at lower temperatures, the vascular contraction 
induced, more than makes up for the covering lost, and is 
consequently followed by a rise of internal body tempera- 
ture; which, like the vascular contraction, is greater as the 
cold is more considerable. Above 70* F. the amount of perspi- 
ration varies with the degree of heat and so far compensates, 
by evaporation, for the differences of temperature as to nudh- 
tain the body at a nearly uniform temperature. 

The temperature of air at which sweating b^ns in the 
nude body is not known. It is at about 86* F. when standing 
at rest. 



ON THE MALAYAN TAPIR, Bhinochoerus sumatranus 
(Gray). By James Mdrie, M.D., F.L.S., F.aS., &c.; 
Lecturer an Comparative Anatomy, Middleeex Hospital: 
fofrmerly Pathologist to ihe Olasg. Boy. Infirmary : Assist, 
Conservator Roy, ColL of Surg. Eng. : and late Prosector to 
ihs Zool. Soc. Land. (Plate viii.) 

1. Preliminary Note and the Outward Features, 

The subject of the present anatomical sketch was a nearly adult 
female animal, which died some eighty hours after its arrival in 
England The stuffed skin and the prepared skeleton are now 
set up among the series of Natural History in the Liverpool 
Museum. The specimen accorded with what goes by the name 
of the Indian, Malay or Sumatran Tapir, often spoken of as the 
Asiatic, in contradistinction to the American speciea Specific 
distinction is based on its being maneless, its hairy coat being 
thin and very short, and the head, neck, fore and hind quarters 
glossy black, with a great bodily girdle of white*, which com- 

^ With such a large mammafl as the Tapir it is not a little remarkable that 
only within the last few years has a new kind been added to soienoe in the 
diaeoYeiy in Panama of a sort combining characters of those already known of 
the Old and New World, bat moreover possessing some striking peonliaxities of 
ita own. 

I may call attention to a lithograph by Mr Wolf (P. Z. S, 1867, PL zlii.) of 
the yomig and adult of this creature, Baird*s Tapir ; the Malay animal has been 
often figured {Tram. Linn. Soe. Vol. zin. &o,), and on aoooont of its hues 
Wagner termed it T. bieolor. 

The literature oonoeming the Tapiridae is ample, especially that deroted to 

the skeleton. Pander and Dalton {VergUieh. Oateol.)t De Blainville {Osteo- 

graphie), others, Wiedemann, (Arehiv. Zool,)^ and Sir E. Home, (Phil, Tran$,) 

Ac. have giTen many delineations of the bones and descriptions thereof, bat by 

far the most lucid account of their osteology is that of Cuvier in his Ossemens 

FogMiles (Tom. n. pt 2, chap. ix. and accompanying Plates). This acute 

obseirer, in giving a r^sum^ of comparison between Indian and American Tapirs, 

was strook by their differences, which however he only considered worthy of 

speeifie distinction, retaining the genus Tapir for both. In a comparatively 

reoent communication on the living Ta|»rine family, Dr J. E. Gray (Notice of a 

new species of American Tapir ^7*. laurillardi)A.e,t P. Z. S, 1867, p. 876, where a 

oopions reference -to anthorities is given), after a careful study of their crania 

divides the group Tapiiid» into three genera — Tapirus, Rhinocharu$t and 

MiatmognathuB. To the first he allots three species, to the second two, and one to 

the last, in all six spedes. Whether these divisions will remain good among 

iiAtoralists the future will show. For my own part I adopt the generic term 

Rkinochofnu as expressive of the cranial difference, which is as trenchantly 

marked from Tapinu., as the former is from the very characteristic Elatmo- 

gnaihu$ Bairdii {QUI, see Yezxill in SilHman's Amer. Joum. Science, July 1867, 

and above-mentioned paper). A living speoimen of this animal has for the first 

9—2 



132 DR MUKIE. 

menced about four inches behind the scapula and extended 
backwards to within a couple of inches of the tail. The white 
colour did not quite meet in the median abdominal line, a longi- 
tudinal darker band some four inches wide here separating the 
right from the left side ; on the hip also the white reached only 
a very little below the great trochanter. The tail is almost as 
broad at the root as the free-part is long ; this gives it a nearly 
equal-sided triangular form. Its total length from the hinder 
edge of the grey colour to the tip is 5'o inches, but the free 
portion or tail proper is but 35 inches in length. The position 
is peculiar, it seeming to be placed much lower than the sacral 
region, which gives a very dumpy or truncate character to the 
buttocks. Of the four toe-nails or hoofs belonging to the 
front foot, (vide Fig. 6, pi. 9), the outermost is the smallest, and 
the third hoof is next in size. The innermost or first corre- 

time been brought to this oormtry alive, and lately was exhibited in the Regents 
Park Gardens (see notice and fig. Field, 7 Oct. 1871). One has a difficulty in 
selecting a specific name for our present eastern form, T, indicut, T. suma- 
traniu, T. malayanut, and T. bicolar haying been respectively used by various 
authorities. 

According to th« law of priority the first mentioned synonyme possibly ought 
to stand, but Gray's T. sumatranut denotes well the limitation of its habitat 
(for a Chinese variety is said to exist), and prevents confusion as to coloration, 
now that the parti-coloured Panama iapir is known. 

It behoves me, in no unkindly way, to take note of a slip of my good and true 
friend Dr Gray. In his paper referred to above he says : '*Guvier states that 
the Malay Tapir was discovered in India by M. Duvaucel," and corrects him 
thereupon. It is true this expression is made use of by Cuvier {Oss. Fost. Art. 
IL p. 156), but these words must be understood in a very modified sense, for 
previously in p. 143 he distinctly mentions that his pupils MM. Diard and 
Duvaucel saw the animal in question at Barkpoor near Calcutta, it having been 
transported to the Governor-general the Marquis of Hastings from Sumatra. 
Ouvier'fl pupils afterwards obtained the animal in the woods of Sumatra, they 
dissected specimens and transmitted the skeleton and skin of a female to the 
Museum d'Histoire Naturelle at Paris. I believe tiie real interpretation to be 
that Duvaucel died, and his MS. transmitted home, confusion arose regarding 
Major Farqnhar*s tapir, described AaiaU Soc, 1816 (Gen. Hardwiok's ColL), and 
what the Frencjmien afterwards saw in Sumatra. 

While thus virtually supporting Dr Gray's separation of the modem Tapirs, 
as most useful in some ways, being based on structural variations quite aa 
notable as in admitted ruminant genera, I confess doing so with a certain 
amount of hesitation ; forasmuch as when the numerous fossil forms of tapirs 
are studied in comparison with the recent ones, it almost behoves us to make 
every individual a genus by itself. In fine, were the fossil remnants of tapir 
sufficiently intact and brought together side by side with the skeletons in our 
museums, I believe a very graduated series, such as to baffle detection of the 
ohanges from one to the other, would result. Nay more, the same might ahnosi 
be said iu allusion to the ancient PdUotherium, Paloplotherium, Lophiodon^ 
Coryphodon, etc. which lead by the easiest steps possible from the tapirs them- 
selves to groups usually denoted relnote. Perfection in classification is the 
naturalist's WiU-o'the- Wisp. I am eontent to use such as is appropriate not- 
with3tanding ito deficiencies, which some cavil at but do not improve. 



THE MALAYAN TAPIE. 133 

sponds very nearly with the third, while the second hoof is con- 
siderably the longest and largest of the four. The second hoof is 
broad and shovel-shaped, the anterior free border being rounded; 
the remaining three hoofs are narrower and more pointed. The 
pad (p) or fatty cushion on the sole of the foot is heart-shaped. 
The middle hoof of the three-toed hind limb projects far beyond 
the outer and iraier (Fig. 14). Like the second of the fore foot 
this, or middle digital hoof, is broad and shovel-shaped. The 
inner and the outer hoofs, equivalent to the second and fourth 
nail-digits, are nearly equal in length, but are not quite half the 
length of the middle hoof, They agree with the outer hoofs of 
the fore foot in being narrowed and sharp at the tips. The sole 
of the hind is narrower than that of the fore foot. 

The following measarements of the body were taken by me : — 

Total length — following the curve of the back 87 inches. 

Height at the shoulder ... ... ... 39 ... 

haunch ... .. ... 40 

Girth round chest (at the anterior margin of 

the white colour) ... ... ... ... 54 ... 

middle of the belly .. ... 60 ... 

Head — from the tip of the snout to the occiput 23 ... 
. . . depth in a vertical Une opposite post- 
angle mandible ... ... ... ... 11*5 ... 

The small elliptical eye is 1 '25 inch in its longest diameter. Its 
exact relative position is four inches from the summit of the head 
and sIk and a half inches from the base of the lower jaw. Between 
the posterior angle of the mouth and the anterior (or inner) angle of 
the eye there is a space of four inches ; and between the posterior 
angle of the eye and the middle of the ear, seven inches. 

Around the angle of the mouth there is a thin edging of 
longish white hairs with a few intermixed but somewhat scat- 
terred long black ones. At the tip of the under surface of the 
proboscis below the nostrils are a few diffused bristle-like hairs, 
easily pulled out, so loosely are their roots attached to the skin. 
Over the proboscis, and in fact distributed here and there over 
the greater part of the front of the head, are longer separate 
black hairs. The hair on the dorsal region of all the feet does 
not quite reach the hoofs, but stops short about a quarter of an 
inch from them, being defined by a sharp line of demarcation. 
In this respect the dorsal hairy covering resembles the plantar 



134 DK MURIE. 

one in terminating abruptly. An injury had befaUen the ea«. 
SO that both were imperfect. The left, which remained in the 
best condition, measured 5 inches long and 3 broad when in the 
natural condition, but when opened it was 4^ inches across. 
The mammary glands were small, the animal having barely 
reached puberty; and just previously having imdergone the pri- 
vations of a long voyage may have conduced or prevented their 
natural enlargement. Two broad, flat and but slightly projecting 
teats were situated in the inguinal region. Each of these had 
from 6 to 8 perforations, which were observed beneath or when 
the skin was removed, but externally or superficially no open- 
ings were apparent. The labia bordering the vulva are well 
pronounced, and there is an indistinct raphe or shallow sulcus 
running between the vulva and the anus. The distance between 
the middle of the vulva and the centre of the anus is 3*2 inches. 
Length of the vulva 1 inch. 

Although the hide of the Suoiatran Tapir is soft and flexible 
compared with that of the Elephant and Rhinoceros, it still retains 
much of the pachydermatoos character. In the present instance 
when being fresh flayed it felt remarkahly stiff and unyielding to the 
knuckles, causing the operation to he a moat laborious and &tiguing 
one to the wrist. Owen's ^ account of the Indian Rhinoceros recurred to 
my mind, inasmuch as the Tapir's hide is veiy much thicker at the 
neck and shoulders, and is even denser and thicker over the iliac region 
and haunch. The fibres of the panniculus camosus muscle are, as 
it were, interblended with the fibrous sub-cellular dermal tissue, so 
that the hide at this part is veiy closely adherent and difiicult to be 
removed; over other parts of the body and limbs, however, while 
tough it was more yielding. 

I had the curiosity to weigh the skin and found it to be 62 lbs., 
a weight equalling that of an ordinary hornless bullock's hide. The 
smaller sized Tapir therefore clearly has the advantage of thickness 
of tegument, and both surpass that of the horse. 

The anus has above a dozen folds around it and these form radii, 
with small wedge-shaped pouches between the elevations. Indeed 
this portion of the perineum has quite a sacculated appearance. 
Hunter' remarks of T. AmericamiB — "The anus is very large, and 
the common skin of the body terminates all at once in the gut^ not 
becoming gradually thinner and thinner till lost in the intestine; 
this termination looks like a cut edge in the common skin and is a 
little scolloped." 

' Trans. Zool. Soc. Vol. iv. p. 86. 

' Eisays and Observations, Vol. ii. p. 167. 



THE MALAYAN TAPIR. 136 

2. The Flayed Carcass, Neck4igament and the Nasal 

Appendage, 

There was little superincumbent or cutaneous fat to pro- 
duce plumpness of contour. Notwithstanding this, by a full 
development of the flesh a certain harmonious configuration of 
outline resulted. What little fatty envelope there existed was 
in a thin layer disposed unequally over the shoulders and loins, 
and this bore very close resemblance to that of a bullock. The 
flesh had a somewhat darker tint than is usually met with in 
the Bovidae, and in this particular rather approached that of 
the Equidae. The hind limb, as low as the knee-joint, when 
compared with the corresponding region of the fore limb to 
the cubit, had the advantage of fulness in flesh, but both were 
powerfully formed. Below the joints spoken of, the limbs were 
nearly equal in thickness, but the fore leg and foot had very 
slightly the greater circumference. 

As Yarrer has described it in the American Tapir, the ligor 
mentwm rmchae is composed of three parta The first and most 
superficial portion is the narrowest and most cordiform. It has 
attachment to the tubercle of the occipital bone, between the 
lateral curved prominences; posteriorly it ends in the inferior 
and lateral portions of the ligamentum nuchae, joining these at 
the last dorsal spine. The two deeper portions stretch between 
the spines of the atlas and the 7th cervial, and then continue 
backwards to the upper lateral edge of the 12th dorsal spine. 
These adpressed portions give off shoots to the several median 
cervical vertebrae, and apparently also dip between the spines of 
the 1st and 2nd, the 2nd and 3rd, and the 3rd and 4th cervical 
vertebrae. 

The entire ligament appeared to be composed of yellow 
elastic tissue. Concerning ite modus operandi, the first portion 
very powerfully flexes the neck and head. The second deeper 
portions act between the vertebrae, and, together with the super- 
ficial division, keep up .the graceful curve of the neck and back. 
The elasticity of this nuchal ligament is extraordinary, rivalling 
that of the Giraffe, of which Quekett* records its contraction 
from 6 to 4 feet on being separated from attachments. 

* Zool Joum, IV. 211. • Trans. Micros, Soe, m. 46. 



136 DR HURIE. 

Among the whole range of mnsctilar anatomy there is no more 
intricate* and withal harmoniously beautiful construction than that 
of the trunk of an Elephant'. Every movement within the radii 
of a sphere is it capable of, and from a soft gentle pressure of power 
it can gradate to a force of immense momentum. The Tapir's pro* 
boscis is nearly identical in pattern, only foreshortened, and its con- 
stituent elements of a finer texture, so that muscular bundles and 
tendons are with more difficulty traced. The construction, whilst 
complex, is nevertheless simple in design, viz. superficially there are 
layers of lengthened, longitudinal, semispiral fibres; deeply fascicu- 
lar bundles set in radii, transversely and obliquely, to the long axis 
of the organ, and partially intercrossing the one with the other, and 
with the superficial elongate ones. I shall add to Eudes-Delongchamp'b* 
account of the myology of the common American Tapir's nasal 
organ, some remarks on that of the Smnatran species, premising re- 
semblance obtains. My commentary to the preceding best follows 
my notes of the dissection. 

1. Superiorly a long thin strip of muscle springs from the 
depression betwixt the nasal bone, its cartilage, and the upper 
inner rim of the orbit. This strip narrows as it proceeds forwards, 
whilst the fibres mingle with those of its fellow of the opposite 
side at the anterior end of the nasal cartilage. Hence they 
run towards the tip of the proboscis where they become more 
tendinous in structure. 

2. There is a very broad sheet of fleshy substance, thin but 
nevertheless strong, which, stretches from the nasal cartilagi- 
nous angle, and supra-orbital prominence forwards radially or 
fan-like upon the flexible snout quite to<its termination. 

3. Some delicate fibres run round the upper surface of the 
orbit and descend in front of the foramina occupying the infra- 
orbital region. 

4. Opposite the said foramina a tenuous slip is sent off, 
which passes to the side of the nasal passage; and the fibres 
composing it join those of the soft musculo-fatty outer wall of 
the proboscis. 

5. Beneath these there exists a broad- and strong muscular 
band, which partly arises from the orbicularis palpebrarum, but 
mainly is derived by a flat tendon, from the infra-orbital angla 
This goes on to the flexible trunk and there spreads itself upon 
the under surface in a longitudinal direction. 

^ 80, to 40,000 mnsctdar fasciculi are affirmed to obtain. 

' Witness Cnvier's and Lanrilland^s elaborate sketches "Myiologie" 

* Mem, de la Soo. Ldnn, de Normatidie, viii. 



THE MALAYAN TAPIR. 137 

6. Another deeper broad sheet with perpendicular fibres 
springs from within, or beneath the hollow of the nares, and 
laterally covers the upper borders of the maxillary and pre- 
maxillary bone. 

7. Furthermore, the narial wall apart from its schneide- 
rian lining is composed of muscular fatty tissue, or a kind of 
areolar looking substance of flesh and glistening tendinous 
fibriUae, the softer padding and minute vessels filling the inter- 
stices so as to give homogeneity. 

The other muscles of the mouth and cheeks will be subse- 
quently referred to along with those of the neck and head. 

Now as to the homology of the above seven divisions the first 
(Py fig. 1) undoubtedly is that spoken of by Cuvier', Owen', Turner', 
and Delongchamps as the special levator, the " L rostii*' of the latter 
author, and of Veterinarians. To my reading equivalent to the pyra- 
midalis nasi of tbe human subject, here however elongated, and with 
a position of ongin corresponding to the alteration of the bony parts. 
The second broad sheet would correspond to the Levator labii superioris 
alaque nasi of man, inc., (Z. Isan) the so-called naso-labial retractor 
in the horse, and the equivalent of Cuvier s* " muscle longitudinal 
anterieur ^ in the Elephant's trunk (a pyramidalis nasi possibly being 
incorporated with it)). Of the third set of fibres in the Tapir these 
seem to agree with the levator anguli oris. The fourth, from posi- 
tion and function, may best be compared with the compressor naris 
or triangularis of human anatomy. In sequence that numbered &ye 
characteristically is to be homologised with the muscle in man, 
termed levator labii superioris propri us (Z. sp)\ its origin, insertion, and 
function being identioed in RhinocJi/oerus as in Homo. Studied with 
respect to the muscles of the trunk of the Elephant, the present, 
third, and fifth muscular series would lead to the inference that they 
are the counterparts of what Cuvier has figured as the '^Muscle 
lateral dirig6 de haut en has et de dedans en dehors* " and '' Muscle 
poeterieur dirig6 de haut en has et de dehors dedans^'* Of the sixth 
muscle in my dissection that known in the human face as the depressor 
alae nasi most coincides with it. In the Elephant it may be in- 
corporated with what is recognised as the inferior longitudinal 
muscle, otherwise it is not differentiated in that animal. Lastly, as 
regards the seventh provisional division, or the deep constituents of 
the narial wall, viz. muscle, tendon and fat, these require much 
minute examination and careful unravelling to elucidate their com- 

1 LeQ(m$, " P. Z. 8, 1831, p. ISS. » P. Z, S. 1850, p. 106. 

4 ** Myologie,** Pis. 274 to 280, and lettered in the latter transverse section 
a, et, ctf a. 

' Ihid. h, b, b, &, PL 280 and PI. 279, fig. 2, No. 5; also more fully delineated 
though unlettered in PI. 276. 

Ibid, e, c, c, c, PI. 280, and well shown unlettered, fig. 2, PL 278. 



^ I 



138 DB BlURIE. 

plex nature. Substantiallj they resemble the radiate and the trans- 
verse fasciculi so ably figured by Ouvier^ I believe them to be, 
in both animals compcured, enlarged homologues of the several 
minute muscles and irregular fibrilke, naso-labials, dca, connected with 
the alar cartilages and upper lip in the human nose. They are the 
same which in part constitute the patulous nostrils of the aberrant 
Saiga*. I have farthermore traced them, with still more remark- 
able changes, in the Sirenia' and Cetacea^j in the latter helping to 
form an immense premaxillary mass of muscle, fat, dec, which 
stretches deeply from the blowhole forwards. 

Finally, I may remark, how varied are the changes of &cial ex- 
pression fix>m the human being to the Tapir, yet the individual 
elements of man's nose and the latter's proboscis bear evidence of 
similitude in their diversity. 

Within the naxes there is a formation of parts of an unusual 
kind. Cuvier^ has curtly limited the topographical relations, 
but Turner' more fully pointed out the structures in question 
also in the American Tapir. His observations show that the 
deep notches situate alongside and partially behind the nasal 
bones are not muscular pits, but curvilinearly afford a nidus for 
the lateral cartilages which form a dilated blind extremity or 
pouch lodged in the said notch. My examination of the same 
species and of the Sumatran animal in the main certify the 
correctness of his statements, though of the latter creature I 
may add a few remarks thereon. In further illustration to 
Turner's diagrams, I append two sketches I made; one, Fig. 8, 
PL X. a semi-dissected fore-segment of the head with the fibro- 
cartilaginous canals and sacs in situ as seen from above, the 
other displaying the interior nasal passages from behind, se- 
parated from the skull with the left convoluted cartilage, gutter 
and sac opened out, the septum nasi being severed vertically. 
(Extending backwards from the anterior soft nares each canal 
lies upon the upward splint of the maxillary, and bending round 
the nasal ends in the bulbous expansion situate betwixt the 
frontal and nasal bones. Thus the two passages together pos- 
sess a heart-shaped outline enclosing the adnate pair of nasals. 
I observed, moreover, that there existed a . kind of everted 

' Loe, eit. d, dfd, d, and e, «, e, e, PL 280, and others. 
« P. Z. 8, 1870, p. 481. 

* ** On the form and stmetnre of the Manatee," a Memoir read before the ZooL 
Soo. 1870, but still nnpubliahed. 

* Vide Joum. of the Linn. Soe. zi. p. 148, and PL 5. 

' '* Lemons WAnaUmie eomparie" * Loe. eit p. 104, figs. 1 and 2. 



THE MALAYAN TAPIR. 139 

tuck, or wliat may be supposed rudimentary subsidiary external 
pouch about the middle of the canal. The entire passage has a 
moist mucous lining, but I did not detect any special glandular 
apparatus. 

The predse fimction of the two canals and double pair of loculi 
I am not in a position to discusa with suitable physiological evidence : 
their homology to me is more apparent Heretofore' I have enun- 
ciated that from position, reLitions, and general structure they cor- 
respond in many ways with the Cetacean spout-hole diverticuli. The 
Tapir's posterior curved canal, as I conceive, is representative of the 
posterior curved nasal pouch of Whales, what I have termed the 
naso-frontal sac {nf&g. 9). The secondary depression, or loculus, men- 
tioned by me above, may be equivalent to a rudimentary maxillary 
sac, lateral pouch of Ceti, or to my so-called &cial occasional division 
of the naso-frontal sac ; its imperfect character giving no very decided 
due, though from what obtains in the ruminant Saiga' preference 
might be given to the former of these. It remains for me, there- 
fore, but to say here that the extra nasal sacs are relatively simple 
in the Tapir, more differentiated in the Saiga, and complex in 
Whales. 

3. Notes an the Visceral Organs. 

The examination of the Thoracic cavity exhibited the 
residual effects of pleurisy, there being fluid exudation, and ad- 
hesions between the parietes and the lungs. The latter were 
congested and exceedingly friable. The disease I conjectured 
had been caught on board ship, very probably resulting from 
the inclement weather of the low latitudes which are visited 
as vessels veer south in following "the great circle sailing." 
In the transport of large mammals from tropical climes, care 
should be taken to provide suitable accommodation, &c., for 
excessive changes of temperature and weather. 

The heart weighed 2 lbs. 3J oz. Diameter at base 7, and 
length from tip to base 7i inches. It corresponded to Owen's' 
description of this organ in the American species, and Poel- 
man's^ account of same in the Sumatran animal, the admeasure- 
ments, however, differing from the latter. Length of the trunk 

^ The Anat. of QlohioeephaUt, ZooL Tratu. read 1867, as yet nnpablished 
by the Society i—On Risso's Grampas, Camb. Joum, of Anat, and Pkyt, v. 
118:— On the White-beaked Bottlenose, Joum. Linn, Soc, v. 141. 

« P. Z. S. 1870, p. 478, flg. 8. 

» P. Z. S. 1881, p. 168. 

* Mem. Acad, Roy. Belif. " Bar le Tapir Indien." 



1,40 



DB MUBIE. 



of the arch of the aorta 3 inches. It then divides into two, of 
which one, the larger, turns to the left forming the descending 
aorta: that to the right is about half the others' calibre, and at 
I inch gives oflF the left subclavian artery. The remainder of 
the branches are in accordance with the Belgian Professor's 
description and figure. 

The Stomach did not quite correspond with the above 
figure, where it has the oesophagus and intestine of about equal 
diameter. The intestine in our female being fully twice the 
diameter of the cardiac portion of the oesophagus. Furthermore, 
the pyloric portion of the stomach curved more towards the 
cardiac end, thus causing the lesser curvature to be smaller 
than depicted. The mucous and muscular coats are, as Poelman 
states, but the glandular portion is deficient at the pyloric and 
cardiac ends, being spread over the middle region. 

Considerable importance has hitherto been attached to the pre- 
sumptive fact of the proportional difference of intestinal length and 
coecum in the Sumatran and American Tapirs. Yarrel*, for instance, 
calculates the length of gut to body in the former as 11 to 1 and in 
the latter 7 to 1, the capacity of the coecum. varying as widely. 
"Without denying diversity in the above respect in the two species, I 
herewith wish to shew that the deduction is manifestly based on im- 
perfect data, and furthermore assert that the alimentary canal's length 
depends quite as much on age, sex, &c, as on mere specific distinction. 
I have compiled the undernoted tabular view to demonstrate this 
point, Poelman's French being converted into English measurement. 



Length, small intestine 
great intestine 
of the coecum 



*i 



)t 



SUMATRAN TAPIB. 


AMERICAN TAPIR 


r 

MmtiE. 


POBLMAN. 


Home. 


MUBIE. 


Turner. 


1 
Owen. 


9 ad. 
ft. in. 


<J ad. 
ft. in. 


(J ad. 
ft. in. 


<? yg. 

ft. m. 


<f yg. 

ft. in. 


(J ad. 
ft. in. 


37 6 
11 2} 
13) 


54 1 
18 114 
-14i 


69 
19 6 
1 — 


28 
4 7 
10 


85 — 
1 1 


45 
9 
1 3 



The valvulflB conniventes and peculiar saccnlate rug» of the colon 
have been sufficiently commented on bj others. I shall only add that 
besides the latter Cetacean character I observed, specially marked in 
T. americanus, structural conformation, smaller but very similar to 
what Dr Cobbold* has described as the ileo-ccecal gland in the giraffe. 



* Zoological Journal, iv. 

« Edinb. Phil. Jour, N,S. ra. 1856. 



THE MALAYAN TAPIR. 141 

Spleen non-adherent to the stomach, but partially attached 
by a portion of the gastro-splenic omentum 1'^ inch from its 
outer wall; the remainder floats free. Weight, 14| ounces. 
It resembled a bullock's in colour and size. Measurements, 
21 inches long, 5 inches broad, and about one-half of an inch 
in thickness. There are no lobules in the Kidneys, as Clift^ 
has observed; on section eight malphigian were counted. The 
right renal organ weighed 13|, the left but 13 ounces. Latter 
^ slightly the shorter and broader of the two. 

In Fig. 7, PI. IX, I give a sketch of the Liver, an organ not 
hitherto pictorially represented (to my knowledge) in the Tapir's 
anatomy. In this Sumatran animal it weighed 6 lbs., and had 
a granular appearance. Its length was considerably greater, 
and its breadth less than Poelman has recorded, and the rela- 
tive magnitude of the individual lobes did not agree with his 
account. In the main Owen's description and Yarrell's short 
notice of the liver of T, Americaniis coincide with what I have 
found both in the same species and T. Stmiatrcmus • 

The left lobe has the advantage of size, the middle next, 
and the right a trifle less than either. Both right and left lobes 
are wanting in marginal clefts: the middle or cystic lobe has 
several. Deep lateral fissures exist between the middle and 
t^ lateral lobes. A tongue-shaped lobus Spigelii is present, and a 

lobule lying upon the Vena cava may correspond with the so- 
called lobus caudatus. 



4. A Review of the Muscles*. 

Fleshy layers of the Trunk, external and internal. 

a. In Cuvier's illustrations the cutaneous layer {Mitscles 
peauciers) is delineated as consisting of three portions; " Tho- 

1 Foot note p. 169, Hunter's Essays and Observ, ii. 

' Appertaining to this sabject, the literature (so far as I am aware) is to be 
found in three publications ; Vrolik issued a memoir {Recherches d'anat. comp. 
8ur le Babyrussa, Amsterdam, 1844), wherein he made a few comparisons of 
its myology and that of the Gnu, the Zebu, and American Tapir. Turner's 
Buccinct paper (as already quoted, 1850, p. 106) contains brief remarks on 
the hyoidean and limb muscles of the latter animal. About the same time 
the posthumus "Recueil" of Gu^ier was issued. It contains 6 figures from 
sketches by himself, evidently of a young (doubtless the foetus glanced at in 
hifl *'Le^n«'-/ animal^ and one figure designed by Laurillard: all seven are 



142 DR MURIE. 

racico-facien" (No. 4), " Portion scapulaire" (5 a), and " Portion 
lat^rale " (5 c). Strictly speaking, this triple division embraces 
what is ordinarily known as the Platysma myoides, and Panni- 
culus camosns. The latter in our animal does not invest the 
.entire trunk as in the genera Sus and Hyrax, and even is more 
limited than in Asinvs and Elephas, though agreeing with 
Equxts Zebra. It has a long pyriform shape athwart the side 
of the chest, from the broad end of which a dense aponeurotic 
fascia runs downwards into the hollow of the flank, and upon 
the side of the knee-joint (Pc^) enwrapping the muscles thereon \ 
Above fleshy, it is attached {P(f) to the middle of nine of the 
last ribs missing the (10th) hindermost. Anteriorly narrow 
{Pc*), it mingles with the fibres of the latissimus dorsi and 
pectoralis major, terminating by fascia spread over the side of 
the forelimb. This fascia extends beneath the deltoid to the 
cubital region, and furthermore mingles with the superficial 
fascia of the lower limb. 

The greatest power of this muscle seems to reside m the hinder 
portion, which thus would have a more direct influence in bringing 
forward the hind leg than in retracting the fore one. This is broiight 
about by the fibres being thickest and broadest at the flank, and the 
line of force is increased by their somewhat semi-lunar direction and 
strongly fixed insertion into the ribs; they also powerfully support 
the abdominal region, and in some degree raise the hind limb. 
Besides being weaker and narrower, they are ntraighter from the ribs 
to the anterior insertion on the outer aspect of the fore limb; the 
line of action being nearly coincident with the axis of the body. 

The portion of the dermal muscle which represents the 
platysma myoides consists of a third broad fleshy sheet covering 
the masseter and in part the mandible, stopping short at the 
anterior third of the horizontal ramus. Aponeurosis fastens it 
to the zygomatic arch and interblends with the zygomatic and 
orbicularis oris muscles. It is noticeable also that a strong 
roundish tendon of the above zygomatic portion attaches itself 

mainly in outline .(Det Plan<ihe§ de Myologies Plates 820—828). The lettering 
indicating masoleB has been loosely put in by A. Fooillon ; and no descriptive 
text aoeompanies the plates, Cavier's sparse remarks -in his "LeQons'* alone 
being applicable. From the above it foUows that a fair aooonnt of the mos- 
cnlo-tenoinons distribation in the Samatran species is legitimate; whilst I 
shall not lose sight of the labours of preceding anatomists. 

^ Alluded to by Vrolik in T. americanw " Reeherches," cited p. 226 as 
" terueur de lafa$eia lata" 



THE MALAYAN TAPIB. 143 

to the anterior and inferior edge of that bone. This division, 
therefore, corresponds to the extra-superficial layer of the mas- 
seter in some Rodents \ 

The Trapessitts^ (Tz.) has a usual situation, and reaches 
almost from the back of the skull to the 8th dorsal spine. It 
is very fleshy, and the nuchal portion is firmly adherent to 
the rhomboid muscle and strong ligamentum nuchae. Remains 
of a slip apparently of this muscle ran towards the paramas- 
toid; but as this was partially destroyed during the process 
of flaying I speak of it with uncertainty. In the Tapir the 
trapezius fixes the scapula as a hinge in movement of the fore 
limbs; its posterior segment retracting the same. The active 
power of bending the head, inasmuch as this muscle is con- 
cerned, is limited; but this deficiency is amply compensated 
for by the strength of the ligamentum nuchae. 

Bhomboidetis\ single fleshy, narrow from the scapula, and 
coming well up to the middle of the back. Derivation, mid- 
neck upon the ligamentum nuchae, backwards. Implantation 
the scapula, Ailly a couple of inches broad, and opposite the 
triangular spot at vertebral border. It approximates the blade- 
bone to the spine, {md produces tension of it forwards. 

Serraius posticus very long, as in Hyrax\ It may be said 
to be divisible into two^ which would represent an anterior 
and inferior portion, which are nevertheless in the main con- 
tinuous by fascia, though their insertions with relations to the 
ribs are different. Its origin, taken as a single one, is by an 
extensive aponeurosis the whole length of the back, from the 
lumbar fascia as far as the most anterior dorsal vertebrae to the 
splenius. This fascia binds down the long dorsal muscles, and 
ends in pointed, triangular, elongated serrations. The insertion 
of the first portion, or serratus posticus posterior {S,p,p.), is 
by seven triangular fleshy slips, that are aflixed to the posterior 



^ Ab a peeuliftrity, I obflerred above the onier meattu Ji denBe .oartUagmons 
ovoid disk 1) by 2 inches in diameter and about a quarter of an inch thick. 
This lay closely applied to the temporal muscle, but interlarded by additional 
glandalar-looking mnsoolar fibres of a strong character. Owen {Trans. Zool, nr.) 
mentions that a similar buccal cartilage exists in Rhinoceroi indictu, 

* Cnvier's, Dorso nu acnnUeny lettered a, a^, a', fig. 1, PI. 822. 
' Portion du Dorso-trachelien, c^ c», Pis. 820, 822. 

* Mnrie and Mivart, P. Z. S, 1866, p. 835. 

^ LauriUard's, PI. 820, Dorso-costien 10, and Lotnbo-eo$tien 11. 



144 DB MIXRIE. 

margins of the same number of last ribs, reaching about mid- 
way between the sacro-lumbalis at its outer border, and the 
upper border of the panniculus camosus. The insertion of the 
second portion, or serratus posticus superior of man, is by seven 
similar tongues, into the anterior borders of the same number 
of ribs; but the seventh rib from behind has one tongue on 
each side, viz. one from each. Thirteen ribs in all therefore 
have attachments. 

The dorsal part of the Serratus niagnus^ {8. mg,) is im- 
mensely thick and fleshy, but it thins considerably as it passes 
to the chest It is continued tolerably strong forwards on the 
side of the neck, being attached to the 7th to 3rd vertebral 
transverse processes. A portion is inserted in the deep inter- 
space between the serratus posticus anterior and the splenius. 
The scapular attachment is wide and firm, so as to produce 
considerable power of slinging the body of the animal. 

Latissimus dorsi^ (La. d.) short, moderately broad, thick 
and strong in the axillary region. Of a fan shape, it lies on 
the side of the chest, stretching from the 9th rib below, forward 
and upwards, to the edge of the scapula and by fascia to the 
dorsal spines above. Here it is covered by the trapezius. It 
is inserted into the lower edge of the teres minor; both muscles 
proceed in union to the humerus. There they lie lower than 
the twist of the brachialis anticus, are placed above the short 
head of the triceps, and are covered by the coraco-brachialis, 
terminating in the inner tuberosity or bicipital ridge of hume- 
rus. The very powerful humeral slip and tendon must be of 
great assistance in retraction of the limb; whilst the enormous 
dorsi-epitrochlearis derived from it raises the lower segment of 
the limb, and drags it backwards through the continuance of 
fleshy fibre of the main portion of the latissimus. 

Pectoralia major*, a moderate, somewhat quadriform mass 
of coarse fibres, which have a direction crossways from the side 
of the thorax to the arm as far as the elbow. It is fixed to 
the sternum opposite the 3rd rib, and forwards to the point of 
the manubrium. Outwardly it lies upon the surface of the 

1 L e. SeapiUo^ottUn g, PI. 320 and fig. 2, PL 822. 
' Chrand dorsal, i. fig. 1, PI. 822. 

* Cnvier's $Um6-humerien portion stemale j, PL 828, fig. 1 ; not, howeyer, 
Lanrillard^s, fig. PL 820 j. 



THE MALAYAN TAPIR. 145 

muscles of the upper arm, from the shoulder to the internal 
condyle. The p. major is in close relation beneath with the 
p. minor, which latter i§ much the larger. Anteriorly its 
margin joins the cephalo-humeral, and it hides the thoracic 
portion of the stemo-scapular. 

Its very thin fibres prevent its having such power as this muscle 
generally has, but the p. miuor being more develo)>ed in a manner 
takes its place. Its chief action seems to be the approximation of 
the limb to the thorax, with a moderate movement of the limb back- 
wards (flexion) j this is thrown foi*ward, as in walking, &c. As 
distinguiiihed from the preceding, the Pectoralis minor* (Pmi) is, on 
the contrary, unusually large'. It covers the surface of the sternal 
cartilages and ribs from about the 2nd to the 10th. Its fibres are 
finer in grain than that of the p. major, and as they taper towards 
the axilla have a slight twist, so that those from behind outside 
come to be undermost, and those at first in front uppermost, or 
most superficial. 

What Mr Mivart and I recognise as the Stemo-scapular' 
muscle in Hyrax* is in Bhinochoerus well-developed, and with 
a somewhat looped figure. Origin, vertebral angle, scapula 
close to the levator anguli scapulae, it broadens as it approaches 
the shoulder-joint, and is attached by fibrous tissue and fascia 
to the cephalo-humeral and pectoralis minor muscles. Thence 
it diverges towards the side of the thorax, spreading out as it 
fixes itself to the three anterior ribs and fourth sternal cartilage, 
but is not attached to the manubrium. From the curved course 
of this muscle it has a raising and protracting action on the 
limb, or, according to the fixed point, would drag the body 
forwards. At the same time the muscles of opposite sides, 
along with the serratus magnus, slings the body on the pivots 
of the fore limbs. 

The muscle now well known as the Supra-costal in the 
Sumatran Tapir is identical with what obtains in the Zebra, 
QiraflFe, &c. It appears almost as if a continuation of the 
scalenus anticus. It is derived, however, by a flat tendon, 
1^ inch broad, from the Ist rib, and proceeds to the cartilage of 

> Portion costale ttemo humtHen, j^ fig. 2, PI. 823, and j by error,, PI. 820. 

' Vrolik describes the long pectorid fibres (above referred to) in the Baby- 
mssa as P. minor, by some considered as a second part of the P. major. 

» The petit pectoral, costo-ooraooidien, Cuv. /. fig. 1, PL 823. It is the sons 
clavier of Vrolik iu the BabymSBa, p. 221. 

< hoc, eit, p. 338. 

VOL. VI. 10 



146 MR MURIE. 

the 4th and the adjoining portion of sternum. Previous to 
reaching the latter it crosses the rectus abdominis. The internal 
intercostala occupy at least seventeen' rib interspaces, and proba- 
bly eighteen, but the latter is somewhat doubtful Commencing 
about the angle of the ribs the external intercostals stop short 
a little way above the commencement of the sternal cartilages. 

In this as well as many other mammals the Triangularis 
stemi is rather to be compared with the dorsal serrati than the 
figure implied by its name. It extends from the 2nd to the 
7th ribs clearly, and then fibres go on from cartilage to bone 
for the rest of the floating ribs ; although the latter might be 
considered as a continuation of the transversalis abdominis. 
The 1st rib is free from this muscle, as in man, the interspace 
being filled with an extension of the deep cervical fascia^ the 
vessels passing between it and the bones. 

The elastic fascia of the External Oblique^, which connects 
it with the ilium and pubis, is very strong; it occupies the angle, 
bounded by the medial line, the anterior rim of the ilium, and 
the pubic symphysis; the tendinous fascia running down the 
middle line of the latter an inch or two. The fascia on leaving 
the ilium is firmly attached to the conjoined psoas and iliacus as 
they pass out of the abdomen, and then continues more loosely 
fixed to a strong slip of aponeurosis, derived from the internal 
obliqua Muscular fibres extend only from the outer border of 
the broad rectus; the interval between each in the median line 
being occupied by an immensely thick, yellow fibre-elastic sheet. 
There are fourteen serrations into as many of the last ribs; the 
four anterior ones interdigitate with the hindermost five of the 
Serratus magnus. Peculiarities of attachment and trifid semi- 
division were noticeable in the Internal Oblique*. A strong 
tendon from the ilium forms a fan-like sheet superficially for 
some inches; muscular fibres diverge forwards to ribs and 
towards the median line of abdomen. Midway between ilium 
and ribs it separated, and the smaller, stronger portion goes to 
the last and tips of cartilages of last two ribs. The other 
broad flat belly stops short of the rectus, and reaches no further 
back than opposite the crest of ilium, while forwards a wide 

* Costo-abdominien, H. 823, fig. 1, No. 18. 
' Ileo-abdominien, 14. 



THE MALAYAN TAPIR. 147 

interspace exists between its border and the rib cartilages. 
The terminal aponeurotic tendon of the internal oblique nearly 
all crosses the rectus; the anterior portion, which in man passes 
behind, in the Tapir is attached to the border of rectus, and 
reaches forwards to the 13th or 14th rib-angle. The Transver^ 
scUia covers internally 9 or 10 rib-cartilages, and its fibres cross 
the anterior half of abdomen; posteriorly it widens as an apo- 
neurotic fascia. The Rectus abdominis^ is derived by strong 
broad teudon, far back on the symphysis; 5 inches forwards it 
becomes fleshy, and with great breadth to the chest enters, 
whence it narrows extremely, and by flat tendon is fastened to the 
Ist rib. Increment of strength to this muscle in the abdominal 
parietes is gained by an aponeurotic fascial strip, which tra< 
verses the costal cartilages and interspaces from the internal 
oblique. There is no Pfframidcdis, 

Considering the actions of the abdominal muscles, the rectus 
appears to be the main central support of the viscera, inasmuch as 
the fibree are thickest, and it is also remarkably broad. The one 
outer half of the internal oblique drags the ribs backwards, while 
its abdominal portion is the great contractor of the outer end of 
the lateral abdominal walls. The external oblique acts reversely to 
the latter, or rather intersects it diagonally, but although slanting in 
direction, it drags back the ribs and also supports the viscera. - The 
muscular fibres of the transversalis being short, it acts mainly on the 
forward half of the abdominal parietes. 

The Psoas parvus* arises by a short tendon from the last 
three ribs at their vertebral junction, and afterwards has a con- 
siderably sized belly, upon the surface of which a flattened 
tendon commences about opposite the second lumbar vertebra. 
This terminates round and strong at the brim of the pelvis, 
midway between the ilium and acetabulum. The Psoas mag^ 
■nus is very broad and arises from the same point outside of the 
above. It covers the transverse processes, also the side of the 
vertebrae, and joins the iliacus. It has a super Hcial tendon half 
its length. The Iliacus is also very broad and fleshy, it has 
origin on the surface of the ilium and along with the psoas 
magnus is inserted into the femur. 



^ SUmo-pubieTU, 15. 

* Pioas et Iliaque rhini, h + i, fig. 1, PI* 328. 



10—2 



148 DR MURIEL 

Sacro-lurnbalis (s.t) comparatively broad and rather strong*, 
as is the Longisaimua dorsi*. Large and fleshy, the Spinalis 
dorsi^ goes as far forward as the spine of the fourth dorsal 
vertebra. Semi-spinalis dorai is uncommonly voluminous, so 
that at first sight it might be taken for part of the Spiimlis 
dorsi. It arises in a V-shaped manner, from the eighth large 
spine, filling the interspace of the fifth and sixth where the 
spinalis dorsi narrows, and then becoming much thicker and 
fleshy, tapers at its anterior end into the foremost dorsal or 
cervical spine. The multijidus spince* are thick, filling the 
hollow between the transverse processes and spine of the ver- 
tebrae ; they have a similar origin and insertion as in man. 
Five interspincdea exist between the long spines. One belongs 
to the last cervical and four are dorsal. The remainder of the 
dorsal spines have apparently only strong fibrous tissue in their 
interspaces. 

MUSCULO-TENDINOUS PARTS OF THE ANTERIOR LiMB. 

h. As in Hyrax the Svhscapularis'^ (8.) does not cover the 
entire inner surface of the bone. At its insertion the small but 
strong tendon of the coraco-brachialis crosses it playing in a 
bursal groove. It may be remarked of the teres major* (7! md,), 
that besides bringing the scapula and humerus in nearer ap- 
proach, it, along with the subscapularis, forms a point d'appui for 
the latissimus dorsi of dorsi epitrochlear to act upon the lower 
limb firom the olecranon process. There is a partly double teres 
minor. The second head by a short strong tendon springs from 
a pit beneath the glenoid cavity : 2 inches from its origin it is 
joined by the first longer head, the two being fixed to the 
tuberosity above the deltoid. The supra spinatus'' {S, sp.) is 
broader than the fossa, and the infra spinaius^ {I. sp.), though 
narrower, is equally strong. Increase of breadth of the shoulder 
muscles is due to the stemo-scapular element. The two former 

1 C. fig. 2, PL 322, •**Myologie." » B. loc, cit. 

' A. loc. cit. figs. 2 and 3. 

* Partly shown in fig. 3, PI. 322, H. 

* ScapulO'trochiniefif n, fig. 2, PI. 323. 

* Scapulo-humerien, 0. 

7 Sm-scapulO'trochiterien, 1, PI. 320, and fig. 2, PI. 828. 

8 Soux-scapulo-troehitenenf m, PI. 820, 



THE MALAYAN TAPIR. 149 

are powerful levers of the fore limb. The first plays as a pulley 
upon the upper extremity of the humerus retracting the head 
of the bone and thus throwing out the leg with force. The 
second, besides this action spoken of, which otherwise could be 
partly subdued or impeded, gains further purchase through the 
prominence, and in consequence has also the duty of rotating 
the limb outwards by the additional fulcrum. 

The Deltoid (/>.), as in the Ungulata and Hyrax, is both 
altered in position and relative size to the muscles of the 
shoulder. It is similarly composed of two long and narrow 
slips. The lowest, longest, and largest portion arises from fascia 
over the infra-scapular muscle, and also from the most inferior 
border, by a very strong aponeurotic tendon, and is inserted 
along with the smaller slip of the muscle, but form the main 
and deepest body of the very strong ligamentous tendon. The 
superior smallest slip lies between the last and the insertion of 
the infra-scapular muscle. It is narrow, short, and somewhat 
flattened, deriving its origin from the inferior anterior third of 
the infra-scapularis. Broadening as it descends it covers consi- 
derably the insertion of the first portion of the deltoid. The 
comparative diminutive volume and position of this muscle 
prevents it from having any very great influence in the move- 
ment of the limb. But it steadies the limb and may counteract 
or aid the cephalo-humeral. 

Dorsi epitrochlear (D. ep.) This triangular muscle^ is entirely 
fleshy, and although only moderately thick is nevertheless won- 
derfully strong by reason of its great breadth at the base, viz. 
between 5 and 6 inches. It is derived from the Latissimus dorsi, 
and distant fully a couple of inches from the shaft of the hume- 
rus. Insertion : the olecranon and inner condyle, its fascia 
is moreover continued down the forearm. Through the long 
lever of the Latissimus dorsi, the dorsi epitrochlear would seem 
also to be an antagonist to the triceps (its scapular head), and 
rotate the limb inwards. 

The Sumatran Tapir agrees with Hyrax Capensis* in pos- 
sessing four bellies to the Triceps^ muscle, besides the last-men- 
tioned. In the latter animal the fourth part is differently in- 

1 t» Cuv. PI. 323, fig. 2. * Loc, cit. p. 340. 

' ReciteU, t, t^, t* Scapulo-oleeranien. 



150 BR MUBIE. 

serted^ namely, into the inner condyle and the olecranon, where- 
as the animal under present consideration has it filling up the 
posterior intercondyloid fossa rather than as a point of origin. 
This extra head in the Tapir can hardly be the anconeus \ 
which is a continuation of the triceps, whereas that in question 
is essentially a quadricipital division sui generis. 

Biceps {B,). Strongly muscular and single-headed". Ten- 
don of origin powerful, fixed to the coracoid process^ its outer 
surface being joined by that of the coraco-brachialis. Inser- 
tion, the inner surface head of radius. The cephalo-humeral 
covers its tendon while lying on the occipital groove. It 
rotates the limb very slightly outwards, besides flexing it. The 
coraco-brachialis would rather rotate it inwards^ but also very 
imperfectly. The brachialis anticus and biceps are powerful 
flexors ; taken together they embrace the humerus in a Y-shaped 
manner. Relatively speaking, the coraco-brachiaUs (C 6.) is of 
considerable volume, and with a single strong tendon of origin 
is derived from the coracoid process in conjunction with the 
tendon of biceps. Opposite the shoulder-joint it spreads out 
into a broadish muscle, which is continued muscularly down the 
inner and front sides of the shaft of humerus, its lower 3rd- 
fourth, the biceps covering a great part of its insertion'. The con- 
joined tendon of Teres major and Lat dorsi reaches higher than 
its upper edge of insertion, but also is fixed behind and below 
the same edge. Like what obtains in many Perissodactyla, the 
Brachialis anticus^ {B.a), has a high post-humeral' derivation and 
radial termination inferior to that of the biceps. The insertion 
of these two occupies nearly the upper third of the bone, and by 
fascia they are continued to the ulna. 

The Supinator longus^ {S. I.) is a riband-like muscle, lying 

1 It doefi not oorrespond ^dth Cavier^s EpieondylO'aibiHfn^Ancon^ exUme^ as 
depicted in fig. 2, PL 823 u, which is strangely placed on the inner aspect of the 
limb. Tomer says the anoonens is wanting or confounded with the triceps, 
he, p. 106. Vide Anconeus epitrochlearit, J. Wood, Yar. Human MyoL P. Boy, 
Soc. 1866—8, and authorities therein quoted. 

' As YroUk mentions. CuTier's scapulo-radien, r. 

* Turner notes in the American species its nearly reaching the inner condyle, 
I. c. p. 106, corroborated by Cuvier, fig. 2, PI. 823, q; and both, along with Yrolik, 
agree as to its single belly. 

* ffumero-eubitiefij s, Guy. 

* Alluded to by Turner and Yrolik, whilst the latter notes as peculiar that in 
the Babyrussa the Lion and Zebu there is but one fascial insertion into radius 
of this muscle and the biceps. 

* ffujnero-siU'radien y v. RecueiL 



THE MALAYAN TAPIB. 151 

closely adherent by fascia to the next two to be described, but 
mainly on the ext. carp, longior. It arises from the highest 
point of the (three) outer supra-condyloid ridge, and winding 
round the forearm upon the surface of ihe two mentioned 
extensors, is inserted by a flat, strong tendinous fascia to the 
inner but anterior prominent edge of the lower end of the 
radius, part of the tendon extending to the styloid process. 
Supinator brevis absent. 

The Extensor carpi radialis longior and brevior^ {E. c. r. L 
and b.) are so interblended in their tendons and muscular fibres, 
that it would be hazardous to say they exist as distinct, but for 
the circumstance that a flattened broadish tendon springs from 
the anterior intercondyloid fossa, and this might represent the 
extensor carp, brevior, while the main origin of the united 
muscle arises broadly by fleshy fibres the whole length of the 
outer condyloid ridge. Together they end about the lower end 
of the anterior surface of the radius in a very broad and strong 
tendon, which passes through the innermost broad groove of the 
wrist, being overlain by the ext ossis raetacarpi poUicis, and 
terminate by tendon and aponeurotic fascia upon the distal end 
of the middle (or 2nd-inner) metacarpal bone ; a small slip of 
tendon, however, is given off to the 2nd (or inner) metatarsal, 
which latter, although it cannot be traced upwards, nevertheless 
may represent the tendon of the ext. carp. rad. brevior. 

The Extensor communis digitorum of the fore limb {E. c. d) 
is large and very powerful. Origin, by a bulky tendon inter- 
mixed with muscular fibres, from the outer condyle, and inter- 
muscular fascia, the latter also connecting it with the outer side 
and front aspect of the head of the radius. It is fleshy to the 
proximal end of the next joint, especially on the inner margin. 
About the middle of the anterior edge of the ulna there is 
derived an additional diminutive slip of muscular fibres which 
joins the larger belly of the muscle above the wrist. The ten- 
don of the extensor com. dig. is very broad, flat and strong, and 
proceeds expanding onwards to the distal end of the metacar- 
pals, where it divides into three divisions", which cover the 

^ Humero-tvU'mita carpien, loc, cit d, apparently single. 
* The ontermoet digit reoeiyee a tendon in Cavier's PI. 322, fig. 1. Epicondylo- 
nu-phalangettien «. 



152 DR MURIE. 

dorsal superficies of the 2nd, 3rd and 4jth digits {i.e. the three 
inner ones) to the proximal ends of the distal phalanges. 

Extensor medii digiti (E.me,d.) and Ex. minimi digiti 
(JEm.d.*'*). Under this heading I group three muscles more or 
less united at their origin. Collectively they arise from the 
external condyle, the inter-muscular fascia, the head of the 
ulna, and the ridge of the uli^^r shaft, nearly its whole length. 
Previous to dissection they seem but one long belly and 
tendon, which latter below at the distal end of the outer meta- 
cai-pal forms a flattened expanse of immensely strong fascia. 
Unravelled, what I term the ext. med. digiti separates about 
the wrist, and with a broadish tendon proceeds to near the outer 
digital fork, where the tendon splits and supplies the two outer- 
most digits (ill. and IV.). The external divisions, or what more 
nearly correspond with ext. min. digiti, have two main tendons 
below; one of these goes solely to the outer (rv.) digit, the 
other proceeds chiefly to the same digit, but moreover partly is 
sent on to the third digit, and has beneath it a cartilaginous 
disk. Above this duplicate, ext. min. dig. resolves itself into 
four constituents. Of these, that denoted as 2 fig. 3, is a dimi- 
nutive musculo-tendinous strip, which adheres close against the 
ex. med. dig. (1) ; a very long delicate tendon represents 3 and 
is adpressed to 4, also a longish tiny but muscular belly. These 
three small diflferentiated parts unite below into one tendon, the 
equivalent of the fourth constituent (marked 5), which respec- 
tively form the dual Ex. min. dig.* 

The Extensor ossis metacarpi pollicis is of considerable 
volume, but not so broad comparatively as in some animals. 
It arises from the interosseous membrane and the adjacent sides 
of the ulna and radius, and is fleshy to within the groove com- 
mon to it and the ext. carp. rad. long, and brevier. It forms 
immediately beneath this a very strong tendon which curves 

^ The above SQbdiyisionB of the oater extensors is snggestive of homology 
with the peronei. Nob. 1 and 5 would represent the peroneus longns and bre- 
vis; No. 4, the peroneas tertius; No. 3, peroneus quartus, and No. 2, probably 
peronens quintus, the distinguishing character of the latter according to Huxley 
being its situation on the front aspect of the bone. Whilst collating the above 
extensors as £. med. and E. min. dig. there is quite a possibility of their being 
considered by others as in part an abnormally inserted Extensor indicis and 
Ext. pollicis et indicis; for variations in which consult Wood's exceUent papers 
in the Jftoy. Soc. Proc. 1866 to 1868. 



> 



THE MALAYAN TAPIB. 153 

inwards and forwards to be inserted into the inner surface of 
the indicial (or inner) metacarpal bone; portion of its aponeu- 
rotic fascia has also an attachment to the Extensor primi, and 
Secundi Intemodit, and the Extensor indicis, as in Hyrax, are all 
wanting. The Ejitensor carpi ulnaris {E,c,u.) is larger and 
stronger than the ext. min. dig. It arises from the ext. condyle 
behind the last, and partially from the olecranon process at the 
styloid process of ulna, where it becomes a single broad tendon 
fixed into the proximal end of the 5th (outer) metacarpal bone. 
This muscle has a deep sulcus in its superficial surface, which 
when ripped up gives it greater breadth and a kind of double 
appearance \ 

The distribution of power of the several fore-limb extensors is as 
follows : the exL carp. radicUis kmg, and hrevis act upon the middle 
(2nd inner) digit, as their name implies. The ext. os, met poll, give 
extension of foot with rotation outwards. The ext. com. dig. is a 
simple extensor of the three innermost digits present, the ext. min. 
dig. of the two outer toes, and the ext. carp, ulnaris of the outer (5 th) 
digit, which it may also' abduct. 

Palmaris longus (P. 7.), may be said to arise broadly from 
the inner surface of the olecranon, and continuing muscular for 
a length of 5 inches, becomes tendinous. This latter is broad 
and flat, and enwraps the conjoined flex. sub. and profundus 
above, but the tendon as it advances downwards narrows, and 
after it passes beneath the pisiform bone is firmly united to 
the outer side of their conjoined tendon. A portion of its 
tendinous fascia is continued onwards to the palm, and crossing 
their surface is lost in the palmar fascia : possibly being con^ 
tinned on to the flexor hrevis mantis. The latter {F. b. m.) 
remarkable muscle, as described by Mr Mivart and myself in 
Hyrax", is very manifest in RhinochoBTus, Its fibres occupy the 
broad palmar fascia, and appear to supply all four toes, being 
one more than in the form compared*. 

^ Apparently applicable to the EpitrocklO'Carpien and CabitO'tut-metacarpien, 
C, C^ of fig. 1, PL S22. 

' Loc. cit. p. 342f fig. 6. 

' JtiBt a barely appreciable line of separation ie recognisable between the 
bellies of the flexor snblimis F. s. d. {Epitrochh-phalanginien, x) ^^^ flexor 
profundus digitorum P.p. d. {Portion du Cubito-sotu-orguien, fi) as they run full 
and fleshy to the wrist, wherefore their tendons are distinct. Those of the 
latter are by far the broader, and four in number, whilst the former supply but 
the three inner toes. Besides the fused condylo-ulnar head, a shorter separate 



154 DR AnXRIE. 

iMmhrxcaUs^ {L,\ 3 in number, and each with a good long 
and thick fleshy bellj. The first and largest arises from the 
palmar surface of the deep united flexor tendon about its middle, 
and proceeding in close connection on the inner side of the 
tendons going to the 4 digit (2 from outer side) is implauted by 
a long thread-like tendon into the middle phalanx of the 4 
digit on its inner border. The second arises between the deep 
tendons supplying the 2 and 3 digits, and is inserted by fascia also 
upon the radial inner side of the 3rd (or 2nd inner) digit. The 
third has a thiner belly than the other two, but is neverthe- 
less a more powerful muscle than either by reason of its greater 
length and stronger tendon. It arises from the palmar surface 
and inner (radial) edge of the deep flexor tendon, but has also a 
strong attachment to the radial (inner) tendon of the flexor 
sublimis. Passing beneath, and to the inner side of this last, 
it ends in a comparatively strong tendon implanted into the 
proximal phalanx of the 2nd (or inner) digit. The 3rd lumbricalis 
would seem to be able to abduct the inner (2nd) digit besides 
flexing it, while the other two analogous muscles are simply 
flexors of the digits. Derived by a short flat tendon from the 
internal condyle, the flexor carpi radialis (F. c. r.)* with a mode- 
rate muscular belly extends half way down the forearm where, 
with a strong cordiform tendon, it continues to the wrist, 
gliding in a deep groove on the bone, and being finally inserted. 
A scant pronator radii teres (P. r. t.) exists', and the flexor carpi 
ulnaris {F. c u.) offers no peculiarity. There is above two 
inches in breadth of a muscle covered by tendon stretching be- 
tween the ulna and radius equivalent to the pronator quadratic 
(P. q,)\ Abductor minimi digiti {Ah. m, d). This, as has been 
noted of some other muscles, resembles the condition extant in 
Hyrax; it is very distinct and strongly muscular. It arises 
thin, broad and fleshy from the distal and outer margin of the 
pisiform bone, where it is adherent to the strong deep palmar 
fascia; increasing in thickness it forms a thickish belly, which 
by fascia and tendon passes outwards, and is inserted into the 

belly springB from tho mid-ulnar shaft and adjoiiung membrane. This repre- 
sentation of a Flexor longns poUicie {P. I. p.) joins the enlarged powerful 
tendon of the profundus at the wrist and loses itself therein. 
^ Not referred to by the previoos writers. 
« EpitrocMo-metacarpieH, 7, fig. 2, PI. 828 " Recueil." 
' Tuner, L c. p. 106. « Mention omitted by authors. 



THE WlLAYAS tapir. 15o 

sesamoid of the 5 (outer) digit. The whole of the InUroaaei^ 
of the fore-foot are highly developed, and each composed of 
coarse strong muscular fibres. There are four pairs (/***), 
viz. 2 to each bone. These are somewhat single at their origin, 
which is from the proximal end of the metacarpals and the deep 
surface of the carpal bone. The inner (radial) one is larger 
than the other and its tendon extends along the side of the 
metacarpal bone to the proximal phalanx, the others are in- 
serted by very strong fascia into the respective sesamoid bones, 
2 to each metacarpal bone'. 

The Neck and the Head-Muscles. 

c. Cephalo-humeral {0. h.). Powerfully muscular and long*. 
Origin partly by muscular tendinous fibres from the outer sur- 
face of the para-mastoid, and chiefly by coarse fibres fi-om the 
outer margin of the lateral process or expansion of the atlas. 
Towards the shoulder it is narrowish but very solid, and lends 
thickness to the neck. As it covers the supra- and infra-scapular 
muscles it expands considerably, becoming thinner and hiding 
the insertion of the aforesaid scapular muscles, as also the 
deltoidii, together with the upper part of the brachialis anticus 
and humeral head of the triceps: insertion upon these last, 
diminishing in muscular fibres until they intermingle with the 
outer fascia of the limb. 

With the limb fixed it must powerfully flex the neck and act 
antagonistically to the ligamentum nuclue. On the other hand, the 

' Neither figured nor described hitherto. 

' There are besides 3 single interosseous musoles superficial to the last {S, t.), 
the Ist (or outermost) is the largest, and along with its fellows arises from the 
under surface of the deep palmar fascia. It is inserted by a long small round 
tendon into the inner side of the middle phalanx of the outer digit. The 2nd 
single interosseous has a similar origin, and is inserted by a similar tendon to 
the Ist on the inner side of the digit. The 3rd and smallest single interosseous 
is more deeply placed between the double interrossei of the 1st and 2nd digits 
(the inner interspace), and ends as the others by tendon, but somewhat shorter 
on the outer ulnar side of the 1st digit. In addition to the 7 interossei tiiere 
is a Tery small still deeper (or donalf) one between the 2 outer digits; this is 
mainly attached to the inner side of the metacarpal of the 4 digit. The 
interossei from their yery great thickness fill the deep hollow of the plantar 
aspect of foot, and together with the immensely thick common flexor tendon, 
flexor breyis manis, and elastic pad, form a solid cushion behind the toes. 

' Lettered in the *'Myologie*' d, as the Trachelo-brachial of G. Cuvier; 
and evidently corresponding to Tumer*8 ** peculiar muscle,*' loe. cit, p. 106, 
andVrolik's masto-humerien, p. 222, met with in the Babyrussa, Gnu, Tapir, &c. 



156 DR MUaiE. 

elasticity of this last, supportiDg the head and neck, forms a point of 
force whereby the cephalo-humeral drags forwards and upwards the 
limb from the shoulder-joint, besides rotating it inwards. 

Two or three long, strong, rounded, cord-like bellies appear 
to compose the Stemo-mastoid. The first portion or belly, 
which is probably the strongest, and is the most superficial 
ventrally, arises by muscular fibres from the manubrium ; it 
terminates in a very strong tendinous fascia several inches 
long, into the posterior ascending border of the mandible. 
This portion appears to correspond to what has been described 
in the Hyrax as the second part or true sterno-mastoid. The 
second distinct belly is smaller than the last, but of a similar 
shapa Its manubrial conjoined origin is deeper and more 
anterior than the first portion, crosses the direction of the latter 
in the neck, and becoming outermost and above. Its terminal 
long, narrow, roundish tendon intermingles with the third por- 
tion of the muscles, and together they are inserted into the 
paramastoid process. The third separate belly corresponds to 
what has been denominated the cephalo-humeral or cleido-mas- 
toid in Hyrax. In the Tapir as it ascends it lies between the 
pectoralis major and brachialis anticus; and, broadening out 
considerably, covers the inner surface of the shoulder-joint, and 
is superficial to the stemo-scapular. It traverses forwards in 
the neck, becoming cord-like, and lying to the outer side of 
the two other portions of the sterno-mastoid ; and being at last 
inserted with the second described portion into the paramastoid, 
its share of the common insertion being mainly muscular. 

The action of the sterno-mastoid apparently corresponds to the 
attachment of each separate belly. The first would flex the head 
through the mandible, and it may be, also, su]»ply the place of the 
deficient platysma myoides. The second would also bend the head 
and neck, but have a less side-movement. The third seems to act 
according to whether the limb or head be the fixed point. If the 
head, then the limb can be dragged forwards, the forearm aided in 
bending, and the whole limb rotated inwards. 

The superficial jugular vein makes its exit beneath pectoralis 
major, goes along the neck between itB second and third portion, 
turns over the surface of the latter, and between it and the first 
portion sends off superior cervical branches, and continues to the 
upper post angle of the mandible, piercing the parotid gland. 



THE MALAYAN TAPIR. 157 

Thin in muscular fibre the Spleniua^ (^0 covers a large 
superficies. Its attachments are the ligamentum nuchsB and 
fascia joining the serratus posticus (anterior). It covers nearly 
the whole of the deep muscles of the neck, forming with them 
a bulging prominence almost separated from the dorsal muscles 
by the manner in which it is bound down, and is inserted 
apparently into the transverse processes of the "six anterior 
cervical vertebrae. The Complexus is also extensive, and forms 
a sheath or cover for the cervical muscles. Origin, by a strong 
short tendon from the outer occipital spine, and by a broad thin 
muscular sheet from the posterior ridge of the temporal bone. 
Narrowing at the hinder end of the neck and becoming wedge- 
shaped it is finally inserted into the transverse processes of the 
seventh or eighth dorsal vertebrae. Its nuchal portion is in close 
relation with the ligamentum nuchae^ where it is very thick, 
whilst the ventral side is much thinner. 

There are sixteen pairs of levator ea costarum; it may be 
seventeen in all, as what has been described as the scalenus 
posticus appears to answer to the most anterior one. 

A broad strong slip departs from the preceding muscle, 
separating from it about the last cervical vertebra, and con- 
tinuing by itself to the transverse process of the atlas; this is 
the so-called levator anguli scapuke, A long muscle rather 
occupies the side of the neck, arising from the transverse pro- 
cesses of cervicals, and becoming broader is inserted into the 
outer edge of the transverse process (lamina) of atlas, a strong 
tendon continues on to the paramastoid process at its base. 
A second muscle of about equal bulk with the above arises from 
the transverse processes of the 4, 3 and 2 cervical, and is in- 
serted into the posterior angle of transverse process of atlas. 

The short deep posterior neck-muscles are each and all fully 
developed ; of these the rectus posticus major has attachments 
to the sides of the spines of the atlas and axis. Smaller by far 
the rectus posticus mitior proceeds to the first vertebra The 
superior oblique has perpendicularly and laterally-placed fibres 
which proceed from the occipital curved line and paramastoid 
to the atlantoid laminae. By far the largest of these nuchal 
muscles is the obliquus inferior, which has ordinary fixed points. 

' 1. 1, Pis. 820, Q22=i Cervico-mattoiden and Dorsotracltelien. 



158 DR MURIE. 

The rectus lateralis is truly ventral in position. With a basi- 
occipital origin the rectus anticus major is continued by strong 
broad tendons inserted into the cervical vertebrae, the second 
to the sixth. The Rectus anticus miTior has insertion into the 
atlas. 

Each Longus coUi is continued within the chest to the third 
dorsal vertebra, and thence the two muscles cover the entire 
under surface of the neck. Tendons of attachment fix them- 
selves to the bodies of the 3 dorsal, 7th, 5th, 4th, 3rd and 2nd 
cervicals, other superficial tendons likewise go to the transverse 
processes of the 2nd to 5th cervical vertebrae. The Scalenus 
anticus is strong and long bellied, tendons of origin spring from 
the transverse processes of the 4th, 5th and 6th cervicals; 
it terminates on the first* rib. 8, medius less bulky, points of 
origin similar but with a broader insertion ; it goes to the 1st 
and 2nd ribs and intercostal space beyond. A diminutive 
quantity of muscular fibres occupies the angle between the last 
cervical and head of 1st rib, and these I take to be the 8. pos* 
ticus. The last is represented longest by Cuvier^, 

The Temporalis (Te) is thin, but of considerable breadth, and 
it reaches quite to the cranial crest. Its insertion lines both 
sides of the coronoid process ; a separate tendinous slip of the 
outer portion proceeds to the alveolar angle. There is no great 
depth of fibre in the Masseter (Ma.), but this is compensated for 
by very considerable breadth. Relatively the Buccinator (Bu.) 
is large, and increment gained in thickness by the development 
of the buccal glands within. This muscle does not descend so 
far as the inferior border of the mandible. 

The broad encirclement of the mouth by the orbicularis 
oris (0. 0.) is marked, and the fibres of the zygomaticus (Z.) 
intermingle with it No clear line of separation existed between 
the external and internal pterygoid muscles\ 

1 PI. 322, fig. 2, Trachelo-eostien, 6, 6«, &*, 

* Tomer observes of the American apeciefi : — ** There is no separate stylo- 
hyoideus, the digastrioos giving some fibres to the os-hyoides. The Tapir also 
possesses a mascle whose fibres (to nse the words of Cuvier| fill a portion of the 
interval of the two comua of the same side. There is a double pair of thyro- 
arytenoid muscles, the upper being partly contindous with the transverse 
aiytenoid muscle, and forming a powerful constrictor of the glottis.** My own 
notes of the neighbouring parts in the Malayan animal run thus: — Each 
digastric (Cuv. PI. 823, fig. 1, 9.) is of considerable size, and its fleshy belly flat 
and moderately thick. There is no positive median tendon, but along the inner 



> 



the malayan tapir. 159 

The Muscles and Tendons of the Hind Quarters. 

dn With origin from the anterior iliac process and patellar 
insertion the Tensor voffince femoris (7! v.f.) has an unusually 
large and strong muscular belly. This is very thick to the 
border next the groin, but thins towards the outer aspect of the 
limb, being there continuous with the gluteus maximus above 
and biceps below. The three, however, are so interblended in 
muscular fibre, which is again overlaid by fascia, as to be with 
difficulty detached from each other. The great strength of the 
tensor vagiose femoris render it a powerfiil extensor of the limb^. 
The Oluteua maximus (&. mx,) is chiefly aponeurotic, it therefore 
seems an extension of the lumbar fascia continued on to the 
spines of the sacral vertebr8e^ Posteriorly it terminates at the 
commencement of the caudal vertebrse, but here thickens and 
lies in a deepish hollow betwixt the posterior border of the 
gluteus medius and anterior border of the biceps. Its fibres 
curve with the hip and are implanted by a strong aponeurotic 
tendon into the third trochanter. The pulley-like action of the 
g. maximus is very much dependent on its being placed in a 
deep groove behind the g. medius, where in a manner it is kept 
in position by the biceps. There is a wonderful depth of flesh 
in the gluteus medius. This muscle (0.m.d.) extends in a wedge* 
shaped manner between the last rib and the ilium ; behind it 
broadens, covering the whole of the dorsal iliac surface and 
overlapping its outer edge; whilst mesially it is fixed to the 
sacral spines, and its attachment reaches the biceps. Its fleshy 
and tendinous insertion is upon the great trochanter besides 
passing down on either side of it. Gluteus minimus, short 
though strong. Attachments, ilium on front of acetabulum, 
but no higher than the obturator foramen. 

side and npper half for a considerable area it is tendinous. Origin, whole of 
the lower surface of the paramastoid ; its insertion overlies the posterior half of 
the mandible partly covering the pterygoid, and with an osseus attachment on 
the middle third of the lower jaw without advancing to the symphysis. The 
mylo hyoidei {Ibid, letter r), from the shape of the digastrioi, do not reach the 
inferior margin of the jaw till within a third of the interspace between the 
symphyses and the angle. The flesh of the mylo-hyoid is thin. Neither of the 
foregoing naso- facial and pterygoidean muscles of the Tapir have been figured 
in the " Planchet de Myologie" but Eudes-Delongchamps partially supplies the 
hiatus in Mem, de la Soc. Linn, de Normandie^ tom. vii. 

^ Named fascia lata (Ileo-fascien) in letterpress Myologie x. in figs. 

* Sacro-femorien and Ilio-irochanterien^Qi. max. and G. med. a, a^. 



160 DB MURIE. 

There is a large and thick obturator extemus^, and the 
0. intemus covers the obturator membrane within and 
reaches well up the bone of the ilium, its tendon passing out of 
the pelvis beneath the gemelli. Gemellus superior^ — small — 
origin, by tendon from the posterior part of the bone at the 
junction of the ilium with ischium, almost opposite the aceta- 
bulum. G, inferior, beneath the last, and with similar in- 
sertion. The quadratus femoris is very little larger than the 
gemelli, and arises posteriorly to the tendon of the obturator 
intemus. All these muscles go as usual to the trochanteric 
fossa. 

Arising superficially upon the upper surface of the con- 
joined psoas and iliacus just within Poupart's ligament, the 
Sartorius* {Sa.) as a narrow band goes downwards and is in- 
serted superficially, but along with the gracilis, over the tuber- 
osity of the tibia. Below the former muscles the sartorius 
occupies the deep hollow betwixt the adductor magnus and 
vastus intemus, partially covering the adductor as it joins the 
gracilis. Its diminutive volume gives it little power as an ad- 
ductor of the limb. It is absent in Hyrax*. The Gradlia^ in 
this instance is a very large thick muscle, and has an origin 
from three-fourths of the posterior line of symphysis ; its fibres 
run outwards and downwards, and are inserted partly by an 
aponeurotic tendon into the inner surface shaft of the tibia 
below the tuberosity. Another portion of the muscle with a 
separate belly diverges about middle posterior of groin, and 
proceeds in a divergent line from last down the leg, terminating 
in a strong round superficial tendon. Action : an inward rota- 
tion of leg from below knee and flexor conjoined with os-calcis 
muscles. 

^The largest of all the vasti is the V. extemus^ which is 
of unusual magnitude (F. e.). The V, intemus {V, i,) presents 
nothing remarkable. Rectus femoris (R.f) single headed, origin 
by a strong tendon from the front of ilium ; insertion patella. 

1 Soils pubio-trochanterien exteme, d. 
■ Portion des Ischii trochanterienst e. 

' PL 823, fig. 1, Ilio-prmbien, t. * Loc, cit p. 847. 

» Shown but as one fiat sheet u, fig. 1, PI. 823, Pubo-jn-Stibien, 
' m and n. Vast. ext. and int., translated as portion du Tri-fimoro-rotulien. 
The expanse of the former is well displayed in Laarillard's PI. 820. 



V- 



THE MALAYAN TAPIB, 161 

The cruretM^ is lai^e and muscular, as distinct firom the above 
muscles as in Hyrax. It covers the entire length of the shaft 
of the femur. The A. magnua {Ad, mg,)*, very large and thick, 
springs fron^ the whole of the pubic symphysis, and femorally is 
fixed on to the lower half of the linea aspera. The femoral 
artery does not perforate its fibres, but lies superficial and dips 
beneath its lower border to reach the popliteal space. What 
corresponds to the A. hrema [Ad. h.) is attached to the brim of 
the pelvis close to the symphysis, and with admixture of tendon 
and flesh is inserted upon the tibial trochanter and linea aspera 
as fiBir as the mid-shaft of the femur. The Pectinem^ {Pe) much 
shorter than the preceding, comes from the lower pelvic brim, 
symphysis to acetabulum, and goes to the trochanteric fossa. 

The Biceps femoria^ is extraordinarily developed and is triple- 
headed; one (£./'), the longest, arising from the spines of three 
anterior caudal vertebrae, and descending to the tuberosity of 
the ischium, it receives a fresh insertion, while the main belly of 
this enormous mass proceeds over the head of the tibia, where 
it is inserted along with the other bellies by a short broad 
tendon, in front of the head of tibia and patella. Another 
portion {B.f?) arises separately from the front of the ischial 
tuberosity by a broadish tendon, which forms a tolerably large 
belly, at first in union with the next, but lower down separated 
slightly and widening out on the side of the leg, joins the 1st 
insertion into the head of the tibia, while part becomes strongly 
tendinous or aponeurotic and runs down the side of the leg. 
The third division {B.J?) seems a continuation of the firat, or 
may be said to be derived from the tuberosity of ischium; it 
joins in a flattened muscular expanse the belly of the second 
one. It furthermore sends another tendon of fascia down the 
leg, which forms a continuation of the tendo achillis below. 
This latter tendon {B,f? fig. 12) is the deepest of the tendons 
composing the tendo achillis, and is joined by the flattened 
fascia tendon of the soleus, the two afterwaids increasing con* 

^ P, 4e Extentum IHo-rotulien, 

* Short and middle addaotors are alone mentioned in the Explanation of 
Cuvier*8 plates, 1, 1>, PL 323, fig. 1, sous-pubo and sous pubifemorien. They do 
not agree in appearance with what I have found obtain in P. sumatrensis, 

* Portion of des Pubo-fimorieni, k. 

* qlsehio-pSronientq sa portion f 4 moraUf -^qaccessoir coccygien, 

VOL. VI. 11 



162 DB MURIE. 

siderably in breadth and thickness, and ultimately splits (vide 
**fig. 12), a portion going to either side of the os-calcis. 

The Semimembranosus^ is of most extraordinary bulk {Sm), 
the muscular mass being several inches in thickness. It is 
double-headed, or at least has different parts of origin, which 
however run into one another from the vast size of the fleshy 
mass. The main origin, in so far as action upon the limb is 
concerned, is that from the whole length of the border of hori- 
zontal ramus of pubis and ischium, and from part of the outer 
side of same. The secondary origin is by a flattened sheet of 
coarser fibres from the dorsal smface, and outer side of the 
caudal vertebrse from opposite the ischium, half-way along to 
the tip of tail. The single belly of these two origins crosses 
downwards and towards the knee, and terminates in a triple 
insertion, the most superficial one by a broad fascia into the 
inner capsule of the knee-joint, below the patella. Another, 
the most delicate and lowest, by a small long round cord-like 
tendon, which comes out of the deep belly of the muscle and 
is implanted in the surface of the inner tuberosity of the tibia, 
dipping somewhat beneath the internal lateral ligament. The 
last and strongest insertion is placed deeper, and is inserted by 
a broad short tendon into the inner condyle of the femur* Its 
bidk must give great power of flexion and rotation inwards of 
the limb, the caudal portion also adding to the length, and 
altering the direction of the leverage, whilst the latter part may 
be the reason why the tail assumes the odd dependent position 
observed in the living animal. With a double origin like the 
last, but not by any means so thick, the semitendinosus {St.) 
springs from the outer surface and tuberosity of ischium, and 
by a thin narrow band fix>m the caudal vertebra, between the 
biceps and semimembranosus^. These ultimately form a long 
rounded belly terminating in a short broad aponeurotic tendon, 
implanted into the shaft of tibia its middle and inner side. 

The Oastrocnemivs* {(fa.), with the usual condyloid origins, 
forms a fleshy calf; a median sulcus almost dividing it into two 
separate halves. These terminate about the middle of the leg, 

^ Ischio-ioiU'tibiefif b. and acoessoir + a. 
■ IschioprStihien^ r. 

* The musole answering this in the **Myologie** is indicated as portion du 
Bi'fimorO'ealcaniefi, and lettered a . 



THE MALAYAN TAPIB. 163 

in a strong body of tendoD, which, with a slight separation of 
fascia, resolves itself into two. The inner one crosses super- 
ficially to the outer side, but can hardly be separated firom the 
other, as they together are inserted into the os-calcis. This 
manner of crossing of these tendons, as well as a similar crossing 
of that of the plantaris and soleus, besides giving superadded 
power of flexion, must also steady the elevating motion of the 
foot, and slightly through the toe-end of the lever, outwards or 
inwards, according to circumstances. 

From its diminutive size and altogether aborted appearance, 
the Soleus' {So.) is still more remarkable. It is little more than 
a mere cord which arises from the inner side of the last on the 
external posterior surface of external condyle. Proceeding 
down the leg parallel with it, and nearly lying beneath, it termi- 
nates at the lower third in the strong flat tendon common to it 
and biceps which is inserted into the os-calcis, the deepest of the 
tendons of tendo achillis. It moreover sends a small slip of ten- 
don to the tendon of the gastrocnemius, and there is an attach- 
ment with the accessory slip of Plantans, as already described. 
This muscle has almost no belly, but is chiefly composed of 
tendon its whole length, a few muscular fibres being seen only 
at its commencement. Turner* remarks, *' the soleus is wanting, 
and the tibialis posticus is wanting also." 

The Plantaris* (Pla.) is peculiar in being a long cordiform 
muscle, which arises from the external condyle at its highest 
part. Traversing between the condyles in the popliteal space, it 
descends the 1^ near the middle line, and about halfway be- 
comes entirely tendinous, and spreading out over the surface of 
the os-calcis, partly attached to either side, it continues into the 
sole of the foot, where it joins the plantar fascia lying superfi- 
cially and strongly adherent to it. At Ihe proximal end of the 
metatarsals it divides into three strong perforate tendons supply- 
ing the three toes present In the back of the leg, below the 
middle, there is an accessory small slip of fascia-like tendon 
which diverges inwards, joining a slip from the soleus, and ter- 

^ Neither figured nor indicated in the Recueil, 

» P. Z. 8. 1860, p. 106. 

* Not distinctly reoogniBable in Gaider^s drawings, although in the explana- 
tion of Platea the letter y is applied to what is supposed to be the Plantaire 
grite (P^mori-ealcanien)* 

11—2 



164 . I>R MURIE. 

minates in a long compound tendon of biceps> Bolens and gas- 
trocnemeus. 

Of moderate bulk the Tibialis anticua (71k.) has a double 
head of origin^ The main inner one from the upper third of 
the tibia's outer surface and the interosseous membrane. The 
second by aponeurotic sheet from the external femoral condyle 
and from the extensor longus digitorum; it joins the first at 
middle of ulna. Joint insertion^ the proximal end of the inner 
dorsal surface of the second metatarsal bone. 

The Extensor Umgus digitorum* {E. I, d) arises by a single 
head firom the middle and inferior surface of the outer condyle. 
Although medium sized in belly, its fleshy fibres reach the 
ankle-joint before forming a thickish tendon. This at the 
middle of the metatarsals divides into three, respectively sup- 
plying the digits (viz. 2nd, 3rd and 4th); the middle one is 
much the broadest. 

The representative of the Extensor longus poUicia^ (E. L _p.) is 
a small slip of fibre given off superficially from the extensor 
longus digitorum about the lower fourth. This runs parallel to, 
or rather to the inner surface, ends over the proximal end of 
the ex. brevis dig.; but its tendon, moreover, goes backwards 
and outwards beneath the tibia before it is inserted into the foot. 

Although seemingly a single muscle* the Extensor brevis 
digitorum {E, b. d,) can be divided into three thin bellies, united, 
however, towards the toes. Origin, the os-calcis the ankle liga- 
ment and bones beneath. Insertion, proximal phalanx of middle 
toe, by a very broad aponeurotic tendon. At this point a small 
tendo-fascial slip goes separately to the inner sides of the outer 
toes. 

A single Peroneus, the P. longus (P. Z.), is all that obtains 
in the limb of the Sumatran Tapir. It is not large. Origin, 
upper three-fourths of the fibula: the tendon thence curves 
round the styloid process, and, passing along the side of the 
foot, is inserted into the external face of the proximal end of 
the proximal phalanx of the outer digit. My note, made whilst 
engaged in the dissection, runs: No other peronei present; 

* Tihio-9u$-tanien, 8. 

' PiriniO'tui'onguiefiy f, fig. 1, PI. 822. 

* Pironii'fut'On^vien, ^, poorly represented by CuYier. 

* The Oalcanio-fuS'Cmguien, fig. 1, PI. 822 ^. 



THE MALAYAN TAPIR 165 

nevertheless I find in Fig. 1, PL 322, of Cuvier*8 own sketch of 
T, Americanus, an indistinct recognition of doable peronei 
opposite the hock. They are lettered as Peroneisous-tarsim e, 
and Peronei-sus-metatarsien €*. 

The Flexor Umgua digitorum^ {F, I d.) arises firom the upper 
three-fourths of the shaft and head of fibula and from the in- 
terosseous membrane; its muscular belly, however, reaches to 
the 08-calcis, and covers the whole of fibula. It there forms 
a strong tendon behind the os-calcis, which winds round to the 
inner side and forms the very strong thick flattened tendon 
and sole. This is joined by the flex. long, hallucis, opposite the 
proximal end of the metatarsal, and the two unitedly passing 
forward about two inches, again subdivide into the three very 
strong thick and broad perforating tendons of the digits, which 
as usual proceed to the proximal ends of last phalanges, spread- 
ing out thinly the entire breadth of the bones. 

The Flexor hngus haUtusi^ (F. I d.) arises by flattened ten- 
dinous fascia from the (posterior tuberosity) styloid process of 
head of fibula, deeper and within, but in union with the flexor 
longus digitorum. It forms a moderate sized belly which crosses 
inwards about middle of tibia, and ends in a strong narrow round 
tendon above the malleolus. This glides in the groove behind 
the malleolus of tibia and crosses again somewhat outwards in a 
similar shallow groove, joining the broad plantar tendon of the 
flex. long. dig. at the proximal end of metatarsal. Of Tapir 
Americanvs Turner {I. c. p. 106) says: " The flexor longus poUicis 
is here, as in all the lower animals, the principal flexor of the 
toes'' — a remark not quite appropriate to the Sumatran species. 

As in the fore-limb there are three lumbricalea* in the hind 
foot, the outer being much the largest. This last {L\ Fig. 12) 
arises between the two outer tendons considerably behind the 
middle one. It has a large belly, which opposite the distal end 
of the metatarsal divides ; and from the muscular fascia of each 
a tendon is derived, the outer one going to the inner side of 
the outermost toe, where it is inserted into the distal end of the 
second phalanx — ^the inner one gives a similar tendon to the 



* TibiO'SOtLS-onguiefij fig. 1, PI. 823, t. 
« Tarsophalangien l\ fig. 1, PI. 828. 

* Not notioed by the observera quoted. 



166 DR MURIE. 

outer side of the middle toe. The middle lumbricalis {V) comes 
from the fork between the second inner deep tendons, but 
chiefly from the middle one, going somewhat round its back or 
deep surface. It is inserted by a small flat tendon into the 
inner side of the proximal end of the proximal phalanx of the 
middle (or 3rd) digit. The innermost lumbricalis (U) springs 
from the inner margin of the inner deep tendon, and is in part 
attached to the inner perforatus tendon of the same digit. It 
sends a delicate tendon of insertion into the inner side of the 
proximal end of the proximal phalanx of the inner (or 2nd) digit 

The muscular fibres of the popliteus {Po, Po*) are very con- 
siderable; including the head of the bone, they extend down 
the posterior surface of the tibia to the middle of its shaft 
Its length probably helps flexion as well as the usual rotation of 
the leg. I did not recognise a Tibialis postictis, agreeing thus 
with Turner's observation in the American Tapir and the draw- 
ings in the Becueil, 

The Interossei^ of the hind-foot are uncommonly like those 
of the fore-foot, both in disposition and general development. 
There are three pairs of double ones (/*, P, /•), attached to the 
metatarsal bones nearly their whole length. The origin of each 
pair is single, terminating in two tendons inserted into each 
side of the proximal end of the proximal phalanges. The middle 
are placed deepest Besides these double muscles there are two 
single ones, smaller in size and placed superficially to the median 
double one, but somewhat overhung by the two lateral double 
ones. The most external (8i^) arises between the inside of the 
outer double one and inner side of middle one. Insertion by a 
tendon to the inner side of outer digit, and into proximal pha- 
lanx, but to the dorsal surface of outer double muscle. The 
internal single one (fift^ is attached in a similar manner outside 
its inner double muscle. 

5. EesuUs of the Present Observations. 

ModeiTL comparative anatomy is suggestive of the fine arts. 
Some revel in creations of their own — airy, fantastic, but always 
eDticing, as ha painting and poetry. Others play the part of the 
sculptor, and by dint of chisel work out the graces of nature her- 

^ Not hitherto oommented on. 



THE UALATAN TAPIB. 167 

Bel£ It has been shown that although the Tapir is wanting in 
the great skin-folds of the Khinoceros, its hide, nevertheless, presents 
similitude in relative thickening at the same points. 

As regards proboscis, it is intermediate between Mephcu and 
RhvnoceroSf shape allying it with the first, dimension and amount 
of prehensility nearing it to the second. With MqutLS it agrees in 
the elongate special levator muscle of the upper lip, while from Sua 
there is a marked separation in there being no terminal cartilagi- 
nous disk. The curious nasal sacs more than remind one of the 
somewhat proboscidean like Saiga amongst the ruminants, and this 

-^ sacculate interior nares carries us to the Cetacean The stomach, gut, 

' and csecum present those curious gradations of structure, leaning to the 
Ruminantia, by tendency to gastric complication, but with stronger 
characteristics shade to its nearer allies among the Perissodactyla. 
But even among the Tapirs themselves shades of division obtain. 
These lead me to suspect that in the visceral organs, as in the 
skeleton, the departures from uniformity show connecting links with 
bygone fossil families. For instance, in the visceral organs of the 
ancient Paleotherium and Hipparion, doubtless intermediance of 
structure existed which would fill up blanks in the living Perisso- 
dactyle group, as indeed the fossil bones afford evidence. 

Among the myological features of interest, as oompai^ed with 
allied forms, may be noted : the restricted area of the panniculus ; the 
multiple division of the biceps femoris and gracilis, and the very low 
insertion of portions of them ; the tendinous nature of the plantaris 
and soleus, their intricate lower divisions and combination with 
gracilis, &c. ; the very full development of interossei and lumbri- 
cales ; the single peroneus ; the splitting up of the outer long digital 

'( extensors of the fore-limb; the presence of flexor brevis manus, 

abductor minimi digiti and pronator quadratus; a belly representing 
flexor brevis pollicis, and the extra division of triceps; the multiplicity 
of the stemo-mastoid ; and the considerable coalescence, fleshy struc- 
ture and elongation of the naso-labial muscles into a proboscis, be- 
sides other peculiarities embodied in the text. Some of these varia- 
tions both agree and differ with the four-toed Hippopotamus, others 
again assimilate and recede from the differently toed families Suidse 
and Equidffi. Characteristic and appreciable changes ensue in the 
fleshy legs and well-padded foot of tiie Elephant ; but, as my paper 
has already assumed goodly proportion, I defer a full comparison 
of our family with others of the oi^ders Perissodactyla, Proboscidea 
and sub-order non-Ruminantia, till a favourable opportunity. 



^ DESCRIPTION OF THE PLATES. 

Plate VIII. 

Fip^. 1. Side Aspect of the Malayan Tapir in the attitude of walking, to 
show in one view the superficial layer of muscles. Drawn from the dissec- 
tion and reduced. 



168 



OR MURIE. 



Pi^j Pe^, and Pc'. PaDnicnlus car- 

nosiis, its seyeral portiona as de- 

cribed. 
L. I. s. a, n. Levator labii superiorig 

alaque nasL 
Py, ryramidaliB nasi. 
Z. s.p. and L. a. o. Levator snperioris 

proprinas and L. anguli ons. 
0. 0. Orbicularis oris. 
Z. Zygomaticos. 
Bu. Buccinator. 
Ma, Maasetor 
^. Splenius. 
Tz, Trapezius. 
S. I, Sacro-lumbalis. 
Lo. d. Longissimus dorsL 
tS.p.p, Senatus posticus posterior. 
Leu cL Latissimus dorsi. 
C?i. Gephalo-humeral. 
St, m, Stemo-mastoid. 
B. a. Brachialis anticus. 
2>. Deltoid. 
T, Triceps and conjoined Dorsi- 

epitrochlear. 



J5. e, r. /. and h. Extensor carpi n^ 

dialis longior and brevior. 
E.o,fn.p, Ext ossis metacarpis 

pollicis. 
E. c. d. Ext. communis digitorum. 
E, me. d, Ext. medii digiti. 
E, m, cL Ext. minimi digitL 
E,c.u, Ext cairpi ulnaris. 
Pa, L Palmaris lougus. 
P. 9, and p. Flexor sublimus and 

profundus. 
O, nue. and G. md. Glutei maximus 

and G. medius. 
T. f>.f. Tensor vaginsQ femoris. 
HP, HP, BiP, Biceps femoris, its 

three divisions. 
S. m. Semimembranosus. 
T.a, Tibialis anticus. 
E.LcL Extensor longus digitorum. 
P. I, Peroneus longus. 
^Retinaculum ankle-joint. 
Ga. Gastrocnemius. 
E.b.d, Extensor brevis digitorum. 
PL h. Flexor longus baJluds. 



Flats IX. 

Different views displaying the myology of the fore-limb; the Liver and 
the front foot 

Fig. 2. Inner muscles of the entire left fore-leg. 

Fig. 3. Semirotate or f view of the outer or extensor tendons and 
muscles of the lower segment of the fore-leg. 

Fig. 4. Distribution of the flexor muscles of the same s^j^ent 

Fig. 5. The Interossei, &C., of the left fore-foot. 

The lettering of these four figures partly agrees the one with the other 
and with fig. 1. 



S. mg, Serratus magnus (portion of). 
S, Subscapularis. 
S. sp. Supra spinatus. 
St, 8, Stemo-scapular. 
C, h, Cephalo-humeraL 

C. b. Coraco-brachialis. 
B, Biceps. 

B. a, Brachialis anticus. 
T, ma. Teres m^or. 

D. ep. Dorsi Epitrochlear. 
T^,T*,T*. Triceps, its three heads. 
S, I. Supinator longus and 
E.cr, I, and h. Extensor carpi ra- 

dialis longior and brevior. 

P. r. t Pronator radii teres. 

Pa. L Palmaris longus. 

P. e. r. Flexor carpi radialis. 

P. e. u. Flex. carp, nlnaria 

P. ». and p. Flex, sublimus and pro- 
fundus. 



P. b. m. Flexor brevis manus. 

E. o, m. p. Extensor ossis metacarpi 
pollicis. 

E, e. d. Extensor communis digito- 
rum. 

E.me.d, Ext medii \ ^ a ^ a k 

_aigin. _ ^ V«l«T««n*Arv 



E m. d Ext minimi f «le™«°**Or 
digiti. ^ 



constituents. 



P. c. u, Ext carpi ulnaris. 

P. 9, d. Flexor sublimus digitorum. 

P,p. d. Flex, profundus digitorum. 

P. l.p. Flexor longus pollicis. 

Z. Lumbricales. 

Ab. m. d. Abductor minimi digitL 

Si^, Si^, Si^. Superior single interos- 

sei (three in ul). 
P, P, P, P. Deep double inter- 

ossei. 



THE MALAYAN TAPIB. 



169 



Fig. 6. Sole of the right fore-foot p. Pad and l n. m. it. digits 
respectively. 

Fig. 7. , The Liver, its posterior or lower surface, 
r. right and I. left lobe. cy. Cystic, q. Quadrate, and sp. Spigelean lobes. 
cL Coronaiy ligament vc. Vena cava. 



Plate X. 

Dissections of the hind-1^ and of the proboscis. 

Fig. 8. Upper view of a fore segment of the face to display the nasal 
canals and sac& The canals are partially opened ; the left eye has been 
scooped out and muscular layer removed on same sida 

N. Nasal bones, r^, Naso frontal sacs, the arrows pointing the course 
of the canals towards the anterior nares. e. Situation of riffht eye, and o. of 
left orbit c. mandibular condyle. X. 1 9. a. n. Levator labii superioris 
alaoue nasi 

Fig. 9. Sketch of a dissection of the anterior nasal passaffes, separated 
from the skull and seen partially from behind but on the rignt side. The 
left canal and sac are opened, the narial septum being sliced transversely 
and vertically. 

an. Anterior nares. W. W.* left and right external walls dragged out 
tj}. septal cartilage, nf. Left naso-frontal sac. / + m. Sinus representing 
either the facial branch of last or maxillary pouch. Arrows lead forwurds: 

Fig. 10. Sole of right hind-foot, n. iil iv., the digits. 

Fig. 1 1. SuperficiS inner view of the entire left lund-log. 

Fig. 12. A deeper view of the groin, &c., same side. 

Fig. 13. Muscles and tendinous distribution of the lower moiety of the 
left hind-limb, seen down-stretched, and from behind. 

Fig. 14. Plantar surface of left hind-foot displaying the interrosseL 

Figures 11 to 14 are lettered as under : — 



R^, Rectus femoris. 

So, Sartorius. 

Pe, Pectineus. 

Ve. and Vi, Vastus extemus and 

intemus. 
Ad.m, Adductor majgnus. 
Ad. b. Adductor brevis. 
Ad. . Gracilia 
tSm. Semimembranosus. 
/SI t Semitendinosus. 
T. a. Tibialis anticus. 
KLd. Extensor lonjgus digitorum. 
E. b. d. Ex. brevis digitorum. 
Oa, Chistrocnemius. 



Bf^, Biceps femoris 3d part inser- 
tion. 

B./.3*. Tendon sent off to join 
soleus, &a 

Pla. Phintaris. 

So. So* Soleus with tendinous 
slip joining gastrocuemius, and ** 
its main terminal division. 

Po. Po.* Popliteus. 

P. I. Poroneus longus. 

P. I. d. Flexor longus digitorum. 

L\ L^J^\ Lumbricales. 

Si\ Si?. Single superficial interossei. 

/*, /*, /'. Beep double interossei. 



V: 



ON THE STERNUM AND VISCERA OF PELL'S OWL, 
Scotopelia peli, Temm. By James Murie, M.D., F.L.S., 
F.G.S., &c. ; Lecturer on Comparative Anatomy at the 
Middlesex Hospital, and late Prosector to the Zoological 
Society, London, (PL XI.) 

Of ike Sternum and SJiovlder-Girdle. 

The reduced views, about two-thirds their natural dimensions, 
which I give of this osteological region, figs. 5 and 6, visually 
express the characters imprinted in the species. As the speci- 
men, however, is of so rare a kind I may describe it in detail, 
to complete its scientific register. 

The extreme length of the sternum from the rostrum to the 
middle xiphoid process is two inches and eight-tenths. The 
greatest breadth is one and nine-tenths of an inch. The per- 
pendicular height, taken in a line stretching from the tip of the 
costal process to the fore part of the keel, gives one-tenth less 
than the above breadth. The keel from end to end, measured 
in a straight line, is equivalent to about 2*4 inches. 

Speaking in a general way the breast-bone is moderately 
strong, but not nearly so powerfully built as in the Falconidse. 
The keel is fair sized and forms a long, regular, curvilinear 
sweep from before backwards and upwards. Its anterior verti- 
cal margiQ possesses a large and deepish bayed notch, sur- 
mounted by a well defined downwardly set rostrum (r.), and 
inferiorly limited by a short obtuse prow-like projection. The 
under surface of the sternum as a whole is quadrangular, a 
third longer than broad ; the fore end has a widish, somewhat 
chevron contour, the hinder end, excluding the processes, is more 
rounded. The lateral edges, nearly parallel, bulge a trifle out- 
wards near the posterior end. Each pectoral plate, lophosteon 
and pleurosteon of Parker ^ has a moderate tilt towards the 
costsB. The ridge which bounds the mid-pectoral muscle is 
short and sharply defined ; the interspace between it and the 

^ I ase freely the ready-coined anatomical terms of my friend^ whose 
laborious researches (Bay Soe. Monograph, 1S68) are a fond of study connected 
with the Tertebrate, shoulder-girdle and sternum. 



STERNUM AND VISCEBA OF PELL's OWL. 171 

ribs being of medium relative breadth, but of fair depth and 
height. Development of costal process (c.p.) is considerable, 
and it pouts well up. Articulations for five sternal ribs exist, 
which latter are long, particularly the hinder ones. There are 
four xiphoid notches, the inner pair shallow but wide, the 
outer pair narrower and deeper. The external xiphoid bar (e.x,) 
is about three quarters of an inch long, middling narrow, and 
tapers to its free end : the internal xiphoid process, also bluntly 
acuminate, barely exceeds 0'2 inch, and does not extend quite 
so far backwards as the middle xiphoid (nu x.) : the latter is 
rounded, the keel terminating in a slightly expanded triangular 
flattened surface. 

The coracoids (or.) throughout are uncommonly stout and 
with great breadth of epicoracoid (e, cr.), slight scooping of the 
angle of which produce a slight ridge upwards on the shaft of 
the bone: the coracoid grooves do not quite meet. The scapular 
end of the coracoid is full and massive. 

Furcula or combined clavicles (cl.) subtend and meet with 
the outline of a gothic arch ; each prascoracoid segment (per,) 
is considerably trifacially enlarged and strong ; from this gra^ 
dually and steadily decreasing almost to a fine spicular rod 
towards the sternal angle, where they join with slight incre- 
ment, but no discernible interclavicle piece^. 

As regards the scapula, its length is about three inches with 
strength in correspondence, the blade being sabre-shaped. The 
glenoideum is large and the acromion, I believe, fair-sized 
Upon the latter point I express myself reservedly, my note 
thereupon being obscurely worded and my original drawing not 
supplying the deficiency. 



The Viscera. 

What remained of the gullet (If inch) was thin-walled and 
nearly uniform in calibre, the muscular fibre being in a dia- 
phanous sheet. Some of the longitudinal mucous rugae are well 
marked below; they are less persistent above, owing to the dila- 

1 In the lithograph the artist has hardly giTen tennity enongh in proporticm 
to the lower moiety of the clavicle, this being most markedly apparent in the 
bones themselTes. 



172 BB MUBIE. 

tability of the lube. Superiorly and within, the uncommonly 
thick glandular wall of the proventriculus ceases abruptly with 
a mitral outline : inferiorly the border is more circular. The 
interior arrangement of the glands is such, that the mucous 
surface to the eye resembles a madrepore coral, being studded 
with minute oscula and puckered divisional . radii. Proventri* 
cular length 1^ inch. In the contracted condition the gizzard 
exteriorly is subcircular, 1^ inch in diameter, and with a central 
tendon as big as a shilling in circumference. Muscular coat 
is evenly distributed and relatively thick : the mucous coat pos- 
sesses complicate folds and a tough epitheleal lining. The small 
gut measures 30, the great gut 2^ inches, or the entire intestinal 
canal is = 32J inches. The lower bowel, as likewise several 
inches of the duodenal loop wherein lays the pancreas, are 
wider, but the average diameter runs about 02 inch. Csecal 
appendages subequal, each about 22 inches in length and pyri^ 
form with moderate dilatation. 

As in most avine forms the liver is bilobed ; these, subequal 
in size, are connected only by a narrow hepatic bridge. But 
the right moiety has a tendency to subsidiary lobulation, inas- 
much as its lower or posterior end, that whereon the gall-bladder 
lies, is smaller and more rounded than the upper or anterior 
portion; and it is partially marked oS from the latter by a shal- f 

low furrow, both ventrally and dorsally. The liver is rather 
over two inches in length, and each half rather under an inch 
and a half in transverse diameter or breadth. The inferior 
vena cava enters the hindermost comer of the right lobe. The 
gall-bladder is egg-shaped, 0*9 by 0*6 in diameter. The bile- 
duct leaves at the post-inferior end. 

All that I obtained of the respiratory apparatus was the 
lower portion of the larynx, the bronchii, and the imperfect 
lungs, a£ depicted in Fig, 3. The latter has the usual spongy 
texture of birds, and, as far as I could make out, are short. .The 
vascular channels are large, from which one would infer vigor- 
ous circulation. One pair of broncho-trachealis (B. t) and one 
pair of stemo-trachealis {8, t) obtain : the former widish. The 
tracheal rings, at least the lower ones, without being bony are 
uncommonly rigid ; their breadth equals the cellular interspace, 
and both are narrow. Each cartilage completes a circle, being 



STEENUM AND VISCERA OF PELL's OWL. 173 

ever so little narrower at the middle, in front and behind. 
The lower larynx or syrinx (la,) has moderate dilatation, and is 
composed of 3 or 4 whole and as many half rings. The antero- 
posterior inner septal division is stoutish and does not rise high. 
The lowermost half ring is much more supple and elastic than 
those above, and it intrudes towards the interior of the cavity so 
as to become valvular in position, and doubtless acts as such in 
vocalization. Semilunar bronchial cartilages, 15 in all, are 
feeble and only half the width of the transparent interstitial 
membrane. 

No special features were detected by me in the heart, which 
after being immersed in spirit, shewed great thickness of the 
left ventricular wall and peculiar longitudinal striations in the 
interior of the aortic arch. 

Each kidney is imperfectly trilobed, the anterior one (a) 
being large, thick and uniform: the middle (m) and the posterior 
(jp) lobules being individually and unitedly very much smaller, 
flatter, and together forming a kind of tailed appendage to the 
former. The ovarium (o) exhibited no enlargement, being alto* 
gether a trifle over 1 inch long ; the ova ranged from a pin's 
head to a small bead in size. The convoluted left oviduct was 
developed as usual. 

ReUuionahip of Scotopdia among the Strigidce, — The very re- 
markable West African form, Peirs Owl, was first referred to by 
Temminck' under SPrix peli. Bonaparte in his " CoDspectus',*' 
places it after Atheney <Scc., and between Ciccaha and Ketv/pa^ as a 
genus by itaelf, Scotopdia. His generic name has in general been 
adopted by succeeding writers: though Prof. Kaup, a long while ago", 
gave grounds for classing it under Ketupa*, The same author like- 
wise arranged it' under the genus Bubo of Cuv., with the following 
as subgenera, viz. ZophostriXy Bubo (Kp.), Ketupa, Urrura and Fseu- 
dotynx. These latter legitimately true genera of other authors, but, 
as he emphatically remarks, 'Hhis is a very grave fault" 

Hartlaub* bas published by far the most explicit description of 
Scotopdia pdi. A specimen is recorded in Duchailla's collection 
and commented on by Cassim^. Lastly, a most excellent figure of 

^ Mus. Lugd. ' Genemm Avium, p. 44. 

* Jardine's Contrib,, 1852, p. 117. A quotation I make Beoond-handed, not 
having been fortonate to obtain the Vol. in question. 

^ In George Gray's recent Hand-List of Birds, Pt. i, Scotopelia forms a 
sabgenus under Ketupa foUowed by Scops, Ao. 

' Monog. of the StrigidiB, Trans. Zool. Soc. it. 2S7. 

* Syst. der Omith. West Africa's, and Cabinis Jour. /. OmitK ni. 358. 
7 Jour. Acad. Nat. Sci, Philadelphia, 1869, p. 52. 



174 DR MURIE. 

the bird, from a living specimen, whicli wbb exhibited' in the Regent'B 
Park Gkirdens, has been given bj Mr John Henrj Gurney in the Ibis^ 
1859, Fl. zv. p. 445 ; and there he notices some slight differences 
from Bonaparte's definition of the species, as well as calls attention 
to its geographical distribation. 

All the preceding authors have formed their judgment on the 
bird solely by its outward characters. I had the opportunity of a 
hasty glance of the sternum and the viscera of the last-mentioned 
specimen, prior to the skin being mounted and transmitted to the 
Norwich Museum. Even with this limited anatomical material 
some additional notion of the bird's affinities are gained. "^ 

As regard the toiU ensemble of the breast bones, the full-sized 
coraooid and scapula, inferiorly tenuous iiircula, and but moderately 
strong sternum, belong, undoubtedly, to the chai*aoters of the Strigi- 
dsa and not the Falconids. In the shoulder-giixUe of all the owls' 
skeletons I have examined, the lower two-thirds of the clavicle tapers 
very considerably to join its fellow without intervention of hypo- 
cleidium ; but Nitzsch' states that in NyctcbUy and Parker' in Athene 
ftoctua^ the clavicles do not meet. In this circumstance Scotopelia 
agrees with the group and not with the above two forms. The 
enlargement and thre&sided condition of each coracoid end of the 
furcula, and widening of epicoraooids, though differing in minor 
detail from certain of the Strigine group, do not afford very pre- 
sumptive evidence of affiliation. Tbe sternum yields appreciable 
characters. In Strix the rostrum is either abortive or absent^ the 
anterior portion of keel is barely produced forwards, the middle and 
internal xiphoid processes are more or less fused together, and the 
external xiphoid spaces and processes relatively very short : points 
trenchantly separating the genus from Scotopelia, Both in Ohts^ ^ 

Brachyotus and Symiimi, rostrum is wanting or very small, the 
front edge of the keel is not emarginate as in Pell's owl, being rather 
straight and perpendicular, with a slight modification in Sy, nebula- 
sum and 0, wlga/ris : the outer and inner xiphoids are slender, 
terminate subequally and relatively long. Nyctcde and Koctua also 
differentiate themselves from our specimen in the internal xiphoid 
bar, being produced considerably beyond the external one. In 
several sterna of Athens examined by me, the rostrum was dimi- 
nutive, in others it was more developed, but in these, unlike S. pelt, 
the fore edge of the keel was straight and vertidd, whilst the 
outer xiphoid is slender and elongate, and the middle xiphoid short 
and narrow. Nyctea {N, nivea) with regard to the shape and develop- 
ment of its processes bears some resemblance to Scotopelia, but 
Eyton's^ plate alone furnishes me data to judge of the posterior 
end of the sternum. In Bvho there is a well pronounced rostrum, 
but this \r carried nearly straight forwards, and there is no anterior 
emargination of keel; the external but slightly surpasses the in^ 

I Prom April 1866 to Oct. 1867. ^*. 

■ Quoted by Kanp, monog, p. 206, ^ 

" Ray Soe. 1868, p. 168. 
* Oiteologia avium^ FL ly. fig. 2. 



STERNUM AND VISCERA OF PELL's OWL. 175 

temal xiphoid, and the middle xiphoid is square rather than ovate. 
In Ketupajlavipea there is a tendency to depression of the prominent 
rostrum, but no inferior emargination, the xiphoids moreover are 
subequal, the middle truncate. The latter character applies to 
K. javoTienns as Ejton has figured \ 

Excepting it may be in the length of the alimentary canal and 
ciecal tubes, the viscera cannot be relied on to determined alliance. 
Their structure in Seotopelia is precisely like the other owls, crop 
being wanting, and therein they differ from the Falconidn save 
Natuilerua /urccthis. 

It results on osteological grounds, that Pell's owl comes nearest 
Eetupa and BtibOj though precisely corresponding to neither. If 
Ketupa is only worthy of subgeneric distinction, as Kaup affirms', 
then Scotopdia likewise must fall under the same ban, but not a 
species of that form. If ornithological genera, however, are ruled by 
anatomical structure. Pell's owl and its nearest ally must rank as 
coequal with Bubo, Notwithstanding, I am satisfied that until a 
fiir more searching scrutiny and analysis of the Strigine osteology is 
worked out, the respective value of the groups, genera and sub* 
genera is but provisional. 



DESCRIPTION OP PLATE XL 

Fig. 1. Renal and reproductiye organs of the female Pell's owl, from 
their ventral aspect f nat size. a. Anterior, m. middle and p. posterior 
lobes of right kidney, o. Ovarium, o, d, oyarian duct or. arfcery and vein. 

Fig. 2. The stomach and lower end of cesophagns opened from the left 
side I nat size, pr, proventriculus, gz, gizzard, py, pyloric orifice. 

Fig. 3. The lower larynx, &c. from in front, of natural dimensions. 
it Stenio-trachealis musde of .the right side, ht Broncho-trachealis. 
Ul Lower larynx, h. Right bronchus. I Portion lung of same side. 

Fig. 4. The alimentaiy canal from the middle of the oesopba^ to 
the vent considerably rednced. oe. oesopfaagns. U tendon of gizzard. 
ccB. C»cal appendagea «p. Spleen. pcL Fancreas. r. Rectum. 

Fig. 6. Tne Sternum and shoulder-girdle, seen from below. Lettering 
as under. 

Fig. & Lateral view of the same. This and above about } nat size. 
9C, scapula, or. coracoid. ecr, epicoracoid. qf>. costal process, r. rostrmn. 
d. clayicle. j^. prsocoracoid. k. keeL ex, external, in, internal, and 
fn». middle xiphoid process. «r. sternal rib& 

» Op. eit. fig. 8. 

' See his final remarks in monog, quoted p. 259. 



ON THE MUSCLES AND NERVES OF A CHIMPANZEE 
(TROGLODYTES NIGER) AND A CYNOCEPHALUS 
ANUBIS. By Frank Champneys, B.A, Brasenose College, 
Oxford. 

Professor Bolleston having kindly provided me with a young Cynooephalns 
Annbis, and subsequently with a female Chimpanzee, I have . dissected the 
greater part of the muscles and nerves of those animals, I have substituted, on 
the advice of Prof. Bolleston, for the name Magot that of Inuus nemestrinus ; 
for this, he informs me, was the correct name of the animal from which Dr 
Church drew his observations. My best thanks are due to Prof. Bolleston for 
assisting me with his valuable advice. I feel also bound to acknowledge, as his 
private property, the ligamentous representative of the long ooraooid in the 
Chimp. 

BOOKS BEFEBBED TO. 

Human Anatomy.— Quain and Sharpey, Ed. 1867. Ellis* Demonstrations, 
Ed. 6. Henle, Handbuch der Anatomie des Menschen. Wilson, AfiatomisVs Vctde 
Mecum, 3rd Ed. Huxley, The Reader, Feb. 13, 1864, and Feb. 20, 1864. Theile, 
Eneyclopidie Anatomique, lu. (French Translation), 1843. Wood, Joum. of 
Anat. and Phys. 1867. Proe. Roy. Soc. June, 1869. 1865, 1868, and 1866 re- 
print. Turner, Joum. of Anat. and Phys. i. 1867 ; Nat. Hist. Rev. Oct. 1864, 
and Trans. R. S. Ed. 1865, 1866, 1867. M4m. de VAcad. de St Pitersbourg, 
1, 1809, and viii. 1859. Luschka, Anat. Mensch. n. 

CoMPABATivB Anatomy.— Yrolik, Recherches d'Anat. sur le ChimpansS, 1841. 
Duvemoy, Archives du Museum d^Hist. Nat. viii. Cuvier, Anatomie C. Vol. i. 
1835 and Planches. Bolleston, Trans. Linn, Soc, June, 1868 ; Forms of Ani- 
mal Life. Parker, Shotdder- Girdle. 1868, pp. 182 — 4, 197, 8. Gegenbaur, Un- 
tersuchungen zur vergleichenden Anatomie, Vol. u. pp. 15, 16. Pagensteoher, Ein 
Vergleich der Muskulatur des Drill mit der des Menschen. Der Zoologische 
Garten, Zeitsehrift filr Beobachtung, Pjlege und Zucht der Tkiere, April and 
May, 1867, Frankfort, p. 128. Notes on a Chimpanzee, MS. Burdach, Beitrag 
zur vergleich. Anat. der Affen. Macalister, Proc. Roy. Irish Acad, x., and Ann. 
and Mag. of Nat. Hist. May, 1871. Wyman, Proc. Boston Soc. of Nat. Hist, 
Nov. 21, 1865, VoL v. Wilder, Boston Joum. of Nat. Hist. 1862, VoL vii. No. 
8, p. 352. Huxley, Med. Times and Gaz tte, 1864, p. 429. Humphry, Joum. 
Anat. and Phys. Vol. i. ,Embleton, Nat. Hist. Rev. April, 1864. Halford, 
^Not like Man, bimaaious and biped, nor yet quadrumanous, but cheiropodous.*'' 
1863. Church, Nat. Hist. Rev. Jan. 1862. Owen, Proc. Zool. Soc. Part i. 
1830— 31. Darwin, Descent of Man. 

Vertebral formula in Chimp. C7, B,,, L4, SC.9 (usually 5 sacral). Dr Em- 
bleton found 10 sacro-coccygeal vertebree. 

The terms "hand** and " foot*' have been used as more convenient than 
*' fore-hand" and "hind-hand." 

The creature had been skinned, to the destruction of nearly all the cutaneous 
nerves. 

The An. was quite young. * Its body was covered with tawny hair, on the 
back it was 24 or 8 inches long, and darker than the rest; on the pectoral 
region it was nearly or quite absent. On the face it was scattered, and there 
were dark bristles on the muzzle and chin. The length of the tail was equal to 
that of the trunk, the proximal half hairy, the distal half bare. 

Muscles. 

The Platysma myoides in the Chimp, was largely developed. It rose 
from the superficial fascitt along a line extending along the external 
half of the external and superior edge of the trapezius. The fibres 



V. 



MUSCLES AND NERVES OF A CHIMPANZEE AND ANUBIS. 177 

ran straight and parallel to be inserted into the anterior part of the 
zygomatic arch, the angle of the mouth and the mental sympliysis, 
becoming continuous with the muscles and fascia in those regions. 

An inch from the symphysis the fibres of the lefl side overlapped 
those of the right. Quain says that when in man there is a decussa- 
tion of jGbres those of the right side overlap those of the left. In 
An. it lay between the two layers of superficial fascia, and arose by 
thin, almost parallel, bands over the region of the Trapezius, Deltoid, 
and upper part of the great Pectoral; ran on each side upwards and 
mesiad, over the clavicle, and over the ramus of the jaw to the sym- 
physis, over which the two sides decussated. A few fibres passed 
over the lower and inner part of the exterior wall of the* cheek- 
pouch, and lost themselves in the superficial fascia covering it\ 

Dermo-humerien (Cuv.), absent in Chimp., was in An. in close 
connection with the skin of the sides and back from the pelvis to the 
axilla, and was coextensive with the Latissimus dorsi, which it 
covered. It was also continued, though with scanty fasciculi, over 
the outer side of the thigh as far as the knee. Its ventral edge was 
the moat muscular, from which it gradually became less muscular 
and more tendinous as it stretched dorsa, losing all its muscular 
fibres opposite the middle line of the back. In the axillaiy region 
its muscular fibres were also few, and in some parts absent; it gave 
a few fibres to the Lat. d. after having previously received a few from 
it. It continued to ascend, and this with greatly increased muscu- 
larity, till it was again gathered up and terminated by a tendou^ 
which was inserted, together with that of the great Pectoral, below 
the head of the Humerus. 

The connection mentioned above seems to be represented occa- 
sionally (Henle and Wood) in man by a connection between Latissi- 
mus dorsi and Pectoralis major; and in the Pigeon by a slip described 
and figured by Rolleston (PI. 2). 

OHiicuiwna (nis in Chimp, was mutilated. 

Levator lahii auperioris, Levator ctngtdi oris present, but not well 
differentiatedL 

Depressor labii vnferioris not distinguishable. 
Zygomaiid were represented by one muscle, half-inch wide, which 
rose from the anterior half of *the zygomatic arch, and from the tem- 
poral fascia above it. Over its mc^ anterior origin a second strip 
rose from the temporal £ascia, but fused with the rest of the muscle 
half an inch below the zygoma. 

StemO'deido-mastoid in Chimp, had a double origin from the 
sternum*. The clavicular tendon, broader than the sternal though 
narrow, arose frt)m the sternal end of the clavicle*. 

^ Cuvier says : " Dans le magot et lea oynoc^phales, il enveloppe T^paole et 
s'^tend en haot on en arri^re dn oou beaucoap plus que dans rhomme.** 

' This double sternal insertion is not found in Man, nor does Yrolik mention 
it in the Ohimp., thouj^ he does (p. 25) remark it in the oran-outan. Guvier 
says nothing about it. 

* Maoalister, Wilder, and Wyman, also found the sternal part larger than the 
davioular part» Duvemoy, in the Gorilla, found the reverse proportion. The 

VOL. VI. • 12 



178 MR CHAMPNEYS. 

AcromuhbaaUar (Vicq. d'Azyr.) or Acromio-trach^ien (Cuvier) 
rose in the Chimp, from the occipital bone on a line with tibe occi- 
pital condyles^, and was inserted into the acromial or external half 
of the clavicle anteriorly to the insertion of the trapezius. In An. 
it was inserted into the exterior third of the clavicle, and rose from 
the ant. side of the transv. pr. of the altas, and by a tendinous slip 
from the occipital. 

Trapezius in Chimp, was as in man. 

Omo-hyoid in Chimp, was as in man*. 

Stemo-hyoid in Chimp, was as in man, and marked nearly in the 
middle with a tendinous portion, as Macalister found it. 

SternO'thyroid in Chtmp. was as in man'. 

ThyrO'hyoid in Chimp, wsa as in man, the anterior or internal 
fibres of the stemo-thyroid being continued into it. 

Great Pectoral in Chimp, rose from the anterior or internal half 
of the clavicle, from the whole length of the sternum, from the car- 
tilages of 8 ribs and the upper of linea semilunaris. Two addi- 
tional strips were differentiated, a superior from the 4th and 5th 
ribs and a fascia stretched yertically between them, an inferior from 
the 5th rib close to the cartilages; these fiised with the rest of the 
muscle opposite the lower border of the axilla\ 

The clavicular portion was not separated by a depression from 
the sternal portion, which depression is mentioned in man in several 
works, e,g, by Yrolik and Ellis. Macalister found no such separation 
in his Chimp. It was inserted as in man. In An. it was as in 
man with two exceptions; Ist^ the clavicular origin extended only 
one-eighth of the length of the clavicle from its sternal end; 2nd, it 
was not distinctly differentiated from the leaser pectoral, as will be 
described. 

The Leaser Pectoral in Chimp, was distinct from the greater Pec- 
toral, rose from the ends of the 1st, 2nd, and 3rd ribs, the aponeu- 
rosis between them, and an aponeurosis from the end of the 4th rib, 
by which its origin extended mesiad as far as the costal cartilages. 
It ended in a round tendon, which was inserted into the capsule of 
the joint, together with the insertion of the supra-spinatus' . 

sternal part is smaller in Cercopithicus, still smaUer in Macacns (rhesns, sinions, 
nemestrinns, and cynomolgus) and Inuns silvanus. VroUk did not find it in 
InnuB. It was the smaller in my An. 

^ It is fonnd in all mammals below Man. Macalister {Pr. I. Ac, %, p. 134> 
says it occurs in one out of sixty hunan subjects. It seems to be correlated 
with a quadrupedal gait. It usually rises from the transverse process of the 
atlas, and sometimes the two following vertebrffi ; and is inserted into the acro- 
mion. It sometimes moreoTer (as in the rabbit) rises from tiie basi-ocoipitaL 

^ The intermediate tendon, Vrolik says, "manque dhez le macaque^ le 
magot, et le babouin." Macalister (p. 848) found it as I did, but feeble. He 
also found it in Inuus and Macacus cynomolgus, and in Cynocephalus porcarius 
and hamadryas. YroUk, he says, did not find it in Inuus and OynocephsJus. 

> Macalister found it marked with a tendinous inscription, which was not 
present in mine. 

* Wood (iV. R, S. 1866) describes in a human subject a separate slip rising 
from the sixth rib, and compares it with '*the so-called chondro-epitroohlear 
muscle of apes and monkeys." 

A Wood found it giving a tendinous slip in several human subjects to the 



MUSCLES AND NERVES OF A CHIMPANZEE AND ANUBIS. 179 

In An. the lesser Pectoral was not a separate muscle, but was 
represented by the deep portion of the great Pectoral. Its coracoid 
insertion was also feeble, and only gained -by the intermediation of 
fascia. (These fibres are described in man by Henle as running 
vertically over the bicipital groove.) It also had an extended inser- 
tion along the exlemal edge of the bicipital groove. 

In Chimp, a distinct fibrous band ran fk>m %he coracoid to the 
sternum between the articulation of the clavicle and the first rib. 
This is said bv RoUeston and others to be the representative of the 
Long Corcbcoid of birds, monotremes, and reptiles. It partially fused 
at its anterior end with the sheath of the subclavius muscle (part of 
the costo-coracoid membrane), but eventually crossed it, and was 
inserted more anteriorly than the origin of the subclavius. It has 
been found by Gegenbaur to contain cartilage oelli^ but there were 
none in my Chimp., though their absence may perhaps be accounted 
for by its adult age. A similar elastic band has been found in a 
Drill by Pagenstecher. 

Svhclamus was in both as in man. 

Scalemts cmticus in Chimp, from the transverse processes of the 
4th to 7th cervical vertebne instead of from the 3rd to 6th as in man. 
It was inserted, as in man, into the " tubercle " of the 1st rib. 

Scalenus posticus did not extend below the 1st rib, as is does in 
nearly all the lower monkeys. In Macalister's Chimp, it did not 
extend below the 2nd rib. In An. the division called posticus 
(proper) was inserted into ribs 3, 4, and 5. 

The supra-pleural fascia (Quain, ii. 894) was well developed in 
Chimp. 

Rectus Thoracis^ absent in Chimp., as also in Macalister's', rose 
in An. from the inferior edge of the 1st rib opposite the insertion 
of the Scalenus anticus and the manubrium stemi, by a short, flat 
tendon, as far as the 3rd rib, where it again became tendinous, and 
was subsequently succeeded by the upper fibres of the Rectus abdo- 
minis*. 

Deltoid in Chimp, arose as in man, and was inserted into the 
deltoid impression on the exterior side of the humerus, the upper 

Cheater tuberosity of the hnmeras. Htunphry (this Journal, i. 366) found its 
insertion extending aeross to the g;Feat tuberosity of the htuneniB in ms Chimp. ; 
Wilder found it iuBerted into the oaracoid on the left, and into the hnmeras on 
the right side of his. 

^ Vamb. Joum. of Anat. and Pkys. May, 1868, p. 893, 4. Henle, p. 96. 

« P. 846. 

* From a simple case like that before us we should call the above muscle a 
prolongation of the rectos abdominis ; bat on yiewing it as it appears in other 
animals («. g. the crocodile), it seems on the whole to be a diyaricated Bm)erior 
portion of the external oblique. It is however also said by Prof, xamer 
(P. R, S. Ed, 1866—7, p. 65) to be closely aUied to the PannicoloB oamosus; 
and he calls it "mosculas stemalis," S. "stemalis brntorom.'' It should be 
mentioned that the inner layer of the greater Pectoral abutted at the upper 
end of its sternal attachment on the tendHnous origin of the ** Beotus thoracis," 
as it expanded to reach the manubrium, and below was continuous by means of 
an aponeurosis with the upper prolongation of the Rectus abdominis mentioned 
above. 

12—2 



180 MR CHAMPNET3. 

end of its insertion being just continuous with the lower and ex- 
ternal part of the insertion of the Peotoralis major (i.e. the part 
rising from the clavicle) ^ In An. it arose from nearly the whole of 
the clavicle. 

Lati89vm,u8 dorsi in Chimp, arose from the spines of the 9th to 
the 13th doi'sal, and all the lumbar vertebree (4) from the supra- 
spinous ligament of that extent^ from more than the outer half of 
the iliac crest, from the 10th to the 13th ribs, was connected with 
the tendons of the dorftal muscles, and with the External Oblique. 
It was inserted into the inner border of the bicipital groove, just 
internal to and alongside of the insertion of the Teres major, a few 
of the tendinous fibres being common to both insertions. 

In An. it differed from that in man in not interdii;itating with 
the External Oblique, but having a long straight aponeurotic origin 
on its ventral aspect about midway between the angles and cartilages 
of the ribs (the interdigitations seem to be replaced by the muscle 
mentioned under Serratus magnus), by failiug to reach the iliac crest 
by nearly two inches, arising instead from an aponeurosis joining 
some of the lower fibres of the External Oblique, and lower down 
from a lumbar aponeurosis, which extended down to the iliac crest'. 
Half-way down it sent a tendon upwards, which joined the lower end 
of the inner division of the Coraco-brachialis^ 

> In Hnmphrv'g, bnt not Macalister's, Chimp., it was oontiauous with the 
Triceps and Brachialis anticus. 

' Dono epitrochlien in Clump, rose from the internal side of the tendon of 
the Latissimus dorsi, about two inches before its insertion, and was inserted in 
the internal condyle of the Humerus. Macalister (p. 844) found it ending in a 
fascia at the middle third of the arm, shorter tiian in most Quadrumana. 
Bergman and Halbertsma describe its anomalous occurrence in man. In mine, 
the muscular fibres extended down the upper two-thirds of Uie arm. With 
regard to the claims of the Dorso-cpitrochlien to be considered a separate muscle, 
and not a part of the Latissimus dorsi, the strongest arguments are derived from 
the innervation. The Dorso-epitroohlien was supplied by a branch which was 
given off by the Musculo-spiral nerve soon after its origin; from the same 
branch a factor was given off which joined the Uhiar nerve just before this sent 
twigs to the inner head of the Triceps. The Latissimus dorsi, on the contrary, 
was supplied separately by a branch of the long Subscapular nerve, which arose 
higher, being given ofiF simultaneously with the Musculo-spiral, but not from it. 

' Yrolik describes the above-mentioned slip as being muscular, and as start- 
ing from the Coraco* brachialis, but wrongly says : ** U va se confondre avec la 
portion interne du triceps.** A glance at his figure (iv. e.) shows that he has 
mistaken the Dorso-epitrochlien (f.) for the inner head of the Triceps. 

The Dorso-epitrochlien is represented (Henle) in man by a constant tendi- 
nous band connecting the long head of the Triceps with the Latissimus dorsi, 
and crossing over the Teres major. This band represents its upper or proximal 
portion, and presents an instiuice of histological substitution. Its dUstal, or 
lower portion, is represented, as it seems to me, in man by the intermuscular 
septum above the internal condyle of the Humerus from which fibres of the 
Pronator radii teres often arise ; these same fibres taking origin in our Chimp, 
(as will be hereafter shown) from the lower part of the tendon of the Dorso- 
epitrochlien. Prof. Bolleston has kindly pointed out to me an account, in a 
book of notes kept in the Biological department of the Oxford Museum, from 
which I quote the following extract : ** Li a child dissected, March, 1862, a dis- 
tinct tendinous band passed down from the broad tendon of the Latissimus 
dorsi to the anterior surface of the long head of the Triceps, which it left at 



MUSCLES AND NERVES OF A CHIMPANZEE AND ANUBIS. 181 

This muscle was well developed in Av. 

Levator am/gvli scapuUje in Chimp, arose as in man from the pos- 
terior transverse processes of the first and second cervical vertebras, 
but differed from that in man in not having a fascicle from the third. 
The superior fascicle split, just before its insertion into the superior 
inch of the posterior border of the Scapula, into several small fiictors. 
The second fuacicle was inserted with the most inferior portion of the 
first fascicle. » 

Ehomboidetis minor in Chimp, was separate from the Bhomboideus 
major', and was as in man. In Ak. the Rhomboidei were feebly 
separated, and the Rhomboideus minor extended up the ligamentum 
nuchse to the occiput. 

Rhomboideus major in Chimp, rose and was inserted as in man, its 
tendon of origin fused with the overlying Rhomboideus minor. A 
curious and complicated fascicle, which must be considered as part of 
the Rhomboideus major, rose, Jirst, fn)m the spines of the 4th and 5th 
dorsal vertebrse, that from the 4t1i being fused with the most poste- 
rior part of the origin of the Rhomboideus major, joining it on its 
deep surface: secondly^ from the spine of the 8th dorsal vertebne. 
From the first origin a muscular slip, I^ in. long, stretchtMi back- 
wards and outwards ; from the second origin a flat tendon, ^ in. long, 
ran forward and outwai'ds, and these met at a right angle opposite 
the 7th dorsal vertebra. From their junction a muscular band 
ran directly outwards like the stem of a Y, the two origins 
forming the two branches. This portion was 3 in. long, and was 
inserted into the most posterior or inferior angle of the Scapida 
with the most inferior part of the Rhomboideus major. At an inch 
from its insertion a few muscular fibres ran directly forwai*d to 
fuse with the posterior part of the Rhomboideus major. An inch 
farther from the insertion the muEcle took a third origin from the 
subjacent fascia, but not from the subjacent ribw. 

No special nervous slip supplied any of the above. 

In An. no such muscle was found, and the Rhomboideus major 
presented no points of note. 

Serratus ma/gnus in Chimp, was stronger than that in man. It 
was formed of three portions. The first, or lowest, arose from the 
1st to the 10th ribs, instead of, as in man, from the 4th to the 8th 
or 9th'. It was inserted into the posterior or inferior angle of the 
Scapula. The second, or middle portion, radiated frx)m its costal 
origin, and its digitations were only slightly marked. It rose from 
the 1st and 2nd ribs and intermediate fescia, instead of, as in man, 

right angles. The mnscnlar slip whioh represents this in the monkey was ' 
quite distinct from the long head of the Triceps in the Ceroopithecns cynosums, 
and in the Hapale penicillata." 

^ Yrolik (p. 18) says the Bhomboids are not differentiated, bnt his was a 
young specimen. Macalister (p. 843) says the same, and states that ibis often 
ocouTS in man. 

^ Macalister found it rising from ribs five to twelve inclnsive. Wilder, from 
ribs five to eleven, and from the fascia covering the Intercostal muscles, by 
slips, of which the lower seven interdigitated with corresponding slips of the 
External oblique. 



182 MB CHAMPNEYS. 

from the 2nd and 3rd ribs. It was inserted into the whole of the 
base of the Scapula^ with the exception of the inch or so oocapied by 
the insertion of portion 1. It was therefore commensui'ate with the 
insertion of the Bhomboideus minor and Levator angoli scapuUe. 
The third portion, which also arose from the Ist rib, and, bj fnsion 
of itH tendon of origin with that of portion 2, also from the 2nd rib, 
was overlapped by portion 2, and was inserted into the anterior or 
superior inch of the posterior or dorsal border (base) of the Scapula 
opposite the insertions of the Levator angulL scapuls. A small 
£iscidle, rising from between the insertions of the Ilio-costalis, Lon- 
gissimus dorsi, and Splenius colli, opposite the 5th cervical vertebra, 
was inserted into portion 3 just an inch from its origin from the 1st 
rib. It seems to come under the category noted by Professor Wood, 
under the head of " occipito-scapular." 

In An. it differed from that in man in having 3 costal attach* 
ments instead of 9. Of these the 1st was inserted into the Ist rib 
in nearly its whole length, the 2nd Into the 2nd rib and fibrous tissue 
between the 2nd and 3rd, the 3rd into the 3rd rib. Of these the 
upper may be said to be again subdivided almost equally by an apo- 
neurosis. It thus extended only as far down as the 3rd instead of 
the 8th rib, as in man. This latter difference, however, was not at 
first sight conspicuous ; for a second muscle, seen on examination to 
belong to a deeper layer, extended down to the interspace between 
the 8th and 9th ribs'. 

SupraspincUiu and Infraspinaius in both were as in man. 

Teres minor in Chimp, differed from that in man, by rising from 
the middle third instead of the anterior two-thirds of the ventral 
edge and adjacent part of the infraspinous fossa of the Scapula. In 
AlN. it was as in man. 

Teres major in Chimp., larger than in man, rose from the pos- 
terior or dorsal ^\ inches of the posterior edge of the Scapula and 
the adjacent part of the infraspinous and subscapular fossse, abutting * 
on, and partially overlapped by, not separated by an inch from the 
long head of the Triceps. It was inserted as in man. There was 
no such connection with the internal head of the Triceps as is 
described by Duvemoy. In An. it arose from the posterior half of 
the Scapula. 

Tricepa and Biceps in both were as in man. 

Coraco-brokchialis in Chimp, was divided into 2 parts, by a cellu- 
lar interval, 1^ in. long, through which the Musculo-cutaneous nerve 

^ This remarkable slip is not found in man, nor was it present in my 
Chimp. It was seen to rise from the angles of the first to the seventh ribs be- 
neath the Serratus magnns in a line between the origins of the Longissimus 
dorsi and Saoro-lombalis, and was attached to ribs four to nine. It seemed to 
be a spedalisation of the external intercostals. 

In an adult An., examined with a view to ascertain the constancy of some of 
the peculiarities noticed in this young animal, the Serratus magnns was found 
to extend to the tenth rib, but the additional slip noticed above was absent. 

Henle mentions a deep layer to the Serratus magnns rising from the first or 
second rib, but fusing with the deep suHace of the muscle. A somewhat similar 
arrangement was found in the Pectoralis major of my Chimp. 



MUSCLES AND NERVES OF A CHIMPANZEE AND ANUBIS. 183 

passed. The iuner of the 2 divisions thus formed was Aised with the 
ooraooid head of the Biceps \ In An. it was soon divided into two 
portions, of which one was inserted into the neck of the Humerus 
on the inner side of the inner edge of the bicipital groove, and cor- 
responded with the 3rd human vaiiety of Wood. The other portion 
was inserted into the middle fifth of the inner side of the Humerus, 
just anteriorly to the origin of the inner head of the Triceps, and in 
a line between that and the Brachialis anticus. It seemed to corr^ 
spend with Wood's Ist variety. It was divided from the Biceps by 
the Musculo-ctttaneouB nerve, which did not pierce the muscle us 
normally in man, or as it did in Chimp. 

Brachialis anticus in both as in man'. 

Subanconetis in Chimp, as in man. 

Anconeus in both as in man. 

Supinator radii longtu in both as in man. 

Extensor carpi radicUis longior in both as in man, except that in 
An. the muscular belly was not differentiated from Ext. c. r. br. In 
Chimp, the remainder of the superficial extensors had, as in man, 
a common origin, and were not differentiated till they had passed one- 
third down the forearm. It will be best to follow the description of 
the arrangement in the human subject^ and to note the differences'. 

Extensor carpi radialis brevior as in man. 

Extensor cowanu/nis digitorum in Chimp, as in man, except that 
the little finger of the right hand received only a slip from the 
tendon going to the ring-finger, just opposite the metacarpo-pbalan- 
gea) articulation ^ 

Extensor minimi digiti in Chimp, was as in man^ In An. it 

^ This arrangement has been noticed in man by Wood {Camb. Joum. of 
Anat. and Phys, 1867, p. 46), and called by Mm Variety 2. There was no third 
Variety of Wood (also known as Goraoo-brachialis brevis vel saperior, and 
Botator humeri). Macalister fonnd a rudiment of it in his Chimp., and he says 
he has found it represented in all Quadrumana. A slip which I have mentioned 
nnder " Dorso-epitrochUen," fonnd by me to be tendinous, by Vrolik to be mus- 
cular, joined the inner cUvision of the Coraoo-brachialis with the Dorso-epitro- 
ohiien. This is described by Wood in Variety 2. 

^ Bolleston found in both arms of a Chimp, a muscular slip parting from the 
upper and outer part of this muscle, and losing itself in the fascia of the fore- 
arm. This is not noticed by Vrolik or Duvemoy, nor did I find it in my 
specimen. 

' Vrolik, while professing to do this (" de mdme que chez Thomme), has de- 
scribed a superficial Extensor Indicis. Now, while it is true that this fascicle 
was as distinct as that of the Extensor carpi radialis brevior, it is also the fact 
that careful dissection will distinguish an Extensor medii digiti, the muscu- 
lar fascicle of which was however covered by the fascicle supplying the Index on 
one side, and by that supplying the little finger (Extensor minimi digiti) on the 
other. 

^ Macalister, Vrolik, and Moore found no tendon to the little finger. Mac- 
alister and Wilder found the tendons not readily divisible : they were readily 
divisible in mine. This slip, as well as one between the ring and middle fingers 
(less well nuurked in the Chimp, than in man), is present in man in addition to 
the proper tendon to the little finger. 

^ This muscle was found by KoUeston in the Chimp. Wood has found in 
the human subject cases of two tendons, two muscles, and an additional tendon 
to the ring finger. Vesalius (i. 258) describes a tendon of this muscle going to 



184 MR CHAMPNEYS. 

differed oonsiderablj from that in man. Rising in common with the 
Extensor commanis digitorum from the external condyle of the 
Humerus, but also from the fsbscia covering the upper end of the 
Radius, it passed down through its proper ring in the annular liga- 
ment, to be inserted into the outer side of the base of the Ist 
phalanx of the little finger, and by a second tendon into the outer 
side of the base of the Ist phalanx of the ring-finger. This arrange- 
ment has been noticed in the human subject by Wood and Vesalius 
(see' note 5 in preceding page). 

Extensor carpi tUnaris in Chimp, as in maii\ In An. it had no 
origin from the ulna. 

Supinator radii hrevia in both as in man. 

Extensor osais fneta,carpi polUcia consisted of two quite separate 
bellies (as its homologue, the Tibialis anticus, did in the leg), having 
the same 2 insertions as the single muscle in man. Of these bellies 
that into the Trapezium, and the prolongation by a tendinous slip 
into the Abductor pollicis (noticed under that muscle), had the more 
superficial orij^in*. In An. it was not double, but had a double in- 
sertion into the Trapezoid bone and the base of the metacarpal bone 
of the thumb, its tendon containing a sesamoid caiiilHge (replaced in 
man by a bursa mucosa) in relation with the quadrupedail habits of 
the animal. 

Extensor prvmi intemodii poUicis was absent in both". In An. 
its absence was more easily detected on account of the singleness of 
the Ext. met. poll. 

Extensor secundi intemodii pollicis in Chimp., as often in man, 
gave a slip to the first phalanx also. It rose from the 2 in. or 2^ in. 
in the middle portion of the Ulna, instead of from its lower half. 
In An. it was as in man. 



join the tendon of the Extensor eonmianis to the ring finger in man. Macalis- 
ter fotmd it in the Chimp., sending a single tendon to the little finger only. 

^ Maoalister found a tendon which he calls **UInaris qninti" prolonged to 
the first phalanx of the little finger ; as in the case of the Peroneus breyis 
(the homologae in the leg of the Extensor carpi olnaris) in my Chimp. This 
is also mentioned in man by Theile and Wood. 

* The muscle is described by Yrolik (p. 20) as two mnsdes which he calls 
** petit Extenseur" and "grand Abdaoteur." He also found an additional tendon 
inserted into a sesamoid bone between the Scaphoid and Trapezium, the two 
others being inserted into the base of the metacarpal bone and the Trapezium 
respectively. Hmnphiy found one tendon inserted into the Scaphoid and meta- 
carpal (which in mine and Maoalister's ended in the Trapezium), the second 
attached to the metacarpal. He says, Yrolik, Wilder, and Wyman found the 
same arrangement as Hmnphry. The muscle has been found in man double, 
and also sending a slip on to the Abductor pollicis, by Wood. {Pr. R. 8, 1866.) 

' Henle says that this muscle sometimes increases its size at the expense of 
the Extensor ossis metacarpi pollicis, which would represent exactly the reverse 
state to that which obtains in the Chimp. The two cases show, however, the 
interdependence of these muscles. BoUeston found the muscle present in 
a Chimp. In my Chimp, two apparently separate muscles which crossed the 
tendons of the radial extensors of the carpus, and seemed to be from their posi- 
tion the Extensores ossis metacarpi and primi intermodii pollicis, respectively, 
were really the two separate divisions of the Extensor ossis metacarpi polliois, 
as was shewn by their insertion. 



MUSCLES AND NERVES OF A CHIMPANZEE AND ANUBIS. 185 

Exiensor Indicts as in man in Chimp.\ In An. it gave a tendon 
to the middle finger, as well as to the Index. 

Flexor carpi tUnaria as in man in Chimp. In An. it rose from 
the internal lateral ligament, as well as from the condyle ; the second 
rose from the Olecranon and the upper end only of the inner and 
anterior border of the Ulna. 

Palmaris longvs in both as in man j it was partly inserted into 
the anterior annular ligament, as in a CniifP. dissected by Bol- 
leston. 

Flexor carpi radialia in Chimp, as in man, except that its tendon 
was not free from muscular fibres till it reached the annular liga- 
ment. In An. it was as in man, but relatively smaller. 

Pronator radii teres in Chimp, was inserted lower down the 
Kadius than in man, and some of its fibres rose fi*om the tendon ci 
the Dorso-epitrochlien '. In An. the second head of origin from the 
ooronoid process of the Ulna was absent. 

Flexor mblirms digitorum in Chimp, as in man, with the follow- 
ing exceptions, which were the same in both hands. The fascicles 
to the different digits were more differentiated proximally than in 
man, and the tendons were much longer. The fascicle to the middle 
fingei* alone, instead of those to the middle and ring fingers, took an 
additional origin from the Kadius'. In An. it had no origin from 

^ It was found by Bolleston and DuTomoy, bat not by Yrolik ; Maoalister 
and Humphry found a second tendon to the middle finger. Wilder found it as 
I did. See Wood for human yarieties. The old dictum that **no ape can 
point" is therefore abundantly disproved. 

* This latter difference, however, was only apparent, for these additional 
fibres rise in man (when they are present) from an intermuscular septum above 
the internal condyle of the Humerus, having the same position as the tendon of 
the Dorso-epitrochlien in the Chimp., and seem to me to furnish the means of 
identifying the above-mentioned intermuscular septum in man as the homo- 
logue of the tendon of the Dorso-epitrochlien. (For further particultu's see 
under the Dorso-epitrochlien. We shall find a parallel instance in the tendon 
of the Glutseus maximum, which furnishes origin to some fibres of the Vastus 
extemus.) The two heads of origin in the Chimp, were divided and innervated 
by the Median nerve, as in man. 

' Macalister found no radial origin. In BoUeston's Chimp., the only fas- 
cicle with a radial origin was that to the Index, I can find no other instance of 
this in man or monkeys : it throws much light on the way in which a muscle 
may, so to say, transfer its origin ; as, for instance, the Flexor hallucis does, the 
homologue of which in the hand rises from the Badius, while it rises from the 
Fibula, the homologue of the Ulna. In my An., as will be seen on referring to 
the description of the Flexor longus hallucis, the latter muscle is seen in the 
process of transference, having half its fibres of origin from the Tibia. By the 
time that man was evolved, the origin of this muscle had, however, quite esta- 
blished itself on the Fibula alone, for Henle remarks that it is very invariable. 
The case plainly put is this : In the same muscular stratum a fascicle rising 
from one or other side is, as a rule, differentiated to a definite digit or insertion ; 
this fascicle, however, by not constantly arriving at the same destination, 
reminds us that the muscular stratum to which it belongs was originally un- 
differentiated, and was capable of a variety of differentiations. The fascicle 
to the little finger arose from the internal condyle of the Humerus, not in 
common with the rest of the Flexor sublimus digitorum (except by a small ten- 
dinous band), but in common with the Flexor carpi ulnaris. MaoaUster says 
Mr Moore found two tendons to the ring finger, none to the little finger^ but 



186 Ma CHAMPNETS. 

the coi^onoid procesB of the Ulna and the oblique line of the Radins. 
Traced upwards from under the annular ligament the tendons deve- 
loped muscular bellies; at middle of the forearm the muscle split 
into tvo portions; the radial side subdivided. One subdivision 
fiised with the Fl. c. r., and ran with it and the Pronator radii teres 
to the internal condyle. The other subdivision fused a little further 
on with a slip (which was a proximally extended representative of a 
common arrangement found in man, and called '^ Fasciculus exilis" by 
Henle^, extending from this muscle to the Flexor longus pollicis or 
Flexor profundus digitorum), which ran from the internal lateral 
ligament and internal condyle to fuse with the Flexor prof, digit, 
with which it had a common origin. The other primaiy portion 
fused first with the FL c. u. and then with the Pal maris longus, with 
both of which it rose from the internal condyle and internal lateral 
ligament. 

Flexor profimdua digitorwm in Chimp, rose (as was also the case 
with the other flexor muscles) from the superior two>thirds instead of 
three- fourths of the Ulna. Its insertion also differed in not including 
a tendon to the Index, which was supplied with one by the Fl. 1. p. ; 
as in all the flexors its tendons were longer and the fascicles more 
differentiated than in man. In both hands the muscular bellies and 
tendons to the middle and ring fingers were more or less closely, 
connected with each other, and with those of Fl. 1. p.' In An. it 
differed from that it man in rising from the upper two-thirds of the 
Badius as well as of the Ulna, and in giving a tendon to the thumb 
(that is, the origin and insertion of the Flexor longus pollicis were 
merged in this muscle). A muscular slip was differentiated to the 
little finger, though it was also attached by connecting &scia to the 
main tendon moving the other fingers. 

Flexor longvs pollicis rose in Chimp, as in man, but had two 
tendons, one to the Index, which was large, while that to the thumb 
was small*. The tendon to the Index pierced the sublimis tendon 

Wilder and Maoalister found four tendons as usual. In both man and Chimp., 
however, the fascicle to the Index is most deeply placed, then that to the middle 
finger, then that to the ring finger. In each case the fascicle to the middle 
finger rising more radially than that to the Index, their tendons cross, that to 
the Index lying the more deeply. In each case the tendons pass under the 
annular ligament in pairs, those to the two middle digits lying superficially. In 
the Chimp, the fascide to the Index was somewhat peculiar ; rising by two dis- 
tinct heads from the same origin, viz. that which was common to ail the super- 
ficial flexor muscles : it developed a tendinous marking on the anterior surface 
of the more internal of these two heads, which only extended for half an inch, 
and gave this portion the appearance of being digastric. Careful inspection, 
however, showed that this tendinous portion took an oblique direction through 
^e fascicle, and appeared again on its deeper or posterior surface. Here it soon 
again disappeared, but an inch above the point of its disappearance a similar 
tendinous marking appeared on the superficial or anterior surface of tiie fascicle^ 
which was continned into the proper tendon. The tendons were inserted into 
the phalanges as in man. 

1 P. 197. 

' Macalister found the muscle not distinct from the Flexor iongus pollicis. 
Wilder fotmd them separate in a Chimp., Duvemoy in a Gorilla. 

' This want of differentiation is sometimes partly retained in man, as 



lIUSCLES AND NERVES OF A CHIMPANZEE AND ANUBIS. 187 

aa if part of the Flexor profrmdns digitorum, ad indeed the whole 
muscle really i». The muscle was so far differentiated into two that 
the muscular fibres, which ended in the tendon going to the thumb, 
extended much farther distally aloDg the common tendon than did 
those which ended in the tendon going to the Index, and these (so 
to speak) would soon become separate in their whole courae. The 
origin of the Fl. 1. p. from the Radius, of the Fl. 1. h. from the 
Fibula (the homologue of the Ulna), has been noticed under the 
Fl. s. d/. In An. it was absent, or rather undifferentiated from the 
FL pr. d. 

FroncUor quadratu8 in both as in man. 

Lumbricalet in Chimp, as in man, except that the 4th arose not 
at all from the tendon of the Fl. pr. d. going to the little finger, but 
from the ulnar side of the tendon going to the ring finger, stretching 
over the tendon and blending at its origin with that of the Lum- 
bricalis of the ring finger. The 2nd, as often in man, rose partly 
from the ulnar side of the tendon of the Fl. pr. d. going to the 
Index. In Ak. they were, on the whole, as in man. 

Falmaris brevis in Chimp, (as far« as could be seen from the shred 
of it which alone was left, the animal having been previously 
skinned) was as in man. 

Abductor poUicis in Chimp, as in man. It was not split into 
slips. Macalister seems to have found it the same. In An. it rose 
by 4 heads, the origin from the annular ligament being subdivided 
into 3 divisions. We shall find several other instances of redupli- 
cation in the muscles of this animal, e,g. in the Iliaous, Psoas, 
Flexor brevis minimi digiti (hand), Extensor brevia digitorum (foot). 
Abductor hallucis, kc. 

Opponeiui pollicis in both as in man*. 

Flexor brevis pollicis in Chimp, as in man*. It was supplied, as 
in mau, by the Median nerve. 

Under tliis muscle we must notice the *^ Interosseus volaris 
primus'^ of Henle. It rose in Chimp, from the radial corner of the 
Os magnum, and from the ligaments covering it, not frx>m the meta- 

obserred by Henle, Wood, and Turner, a muBonlar slip from the Flexor longus 
polUois sending a tendon to join the tendon of the Flexor profandus digitonun 
to the Index. 

^ The tendon to the pollex crossed the other tendons of the Fl. p. d. in 
Hnmphiy's but not Macalister's specimen. Yrolik found no tendon to the 
thumb ; Humphry found the tendon to the thumb rising as a slender tendon 
from the palmar fascia and going to the last phalanx of the thumb in one spe- 
cimen, and in another as a long thin tendon from the ulnar side of the Flexor 
profundus. Wyman found it as Macalister, Wilder found it conjoined with the 
Flexor profundus indieis. In both hands of my specimen there was a good deal 
of tendinous connection at the origin of the tendons with those of the Flexor 
profundus digitorum going to the middle and ring fingers. 

^ Duvemoy describes it in the Chimp, as divided into two portions, one rising 
higher than the other on the annular Ugament, and the longer fascicle inserted 
more externally than the shorter : but this was not the case in my specimen. 
Dr Embleton found it absent in a young Chimp. 

' Macalister found it as I did, but Humphry found the outer portion extend- 
ing beyond the first phalanx. 



188 MR CHAMPNETS. 

carpal directly as in man \ In Ax. the Fl. br. p. seemed to be a^ 
in man. 

Adductor pollioia and Abditctor Tmnimi digUi in both. 

Flexor brevis minvmi digiti in Chimp, arose as in man, but was 
inserted not together -with the Abductor minimi digiti but distally 
to it in the same line, by two tendons, of which the external or 
radial was more distally inserted than internal or ulnar. It also 
gave two tendinous slif>s to the Abductor minimi digiti, with which 
in man it is often fused. In An. it was not marked off from the 
Abductor minimi digiti by the deep branch of the ulnar nerve. The 
head, rising from the annular ligament^ was split into three, placed 
in axial series (thus furnishing another instance of reduplication of 
muscles, such as we have noticed), and the ulnar nerve passed between 
these and the fourth head, which, as part of the muscle in man, rose 
from the tip of the hooked process of the unciform bone. All of 
these heads, except the most radially, which was also the most dis- 
tally placed, of tho.<e rising from the annular ligament, were mure or 
less fused during some part of their course with each other and with 
the Abductor minimi digiti, and were inserted together into the 
base of the first phalanx of the little finger; the most radially placed 
(before mentioned) having a similar but separate insertion. 

Opponens minimi digiti was in both as in man. 

Dorsal irUerossei in Chimp, were 4. Abductors from the axis of 
the middle finger ; the " Abductor indicis " was more largely deve- 
loped than in man. It had a double origin from the index meta- 
carpal, one from the base, the other from about two-thirds of its 
length. The radial artery, as in man, separated the origin of the 
metacarpal of the thumb from that of the metacarpal of the 
Index. It was inserted as in man. The other dorsal Interossei 
were as in man'. 

FcUmcbr interossei six in number in Chimp.", each rose from the 
whole of the metacarpal of the digit into which it was inserted, and 

1 Bolleston found it in his Chimp. Wood mentions its presence in man. 

' Macalister found them all as in man. In Ateles fuliginosus, in which the 
thumb is represented by a radimentaiy metacarpal, he fonnd them thas : " Of 
palmar Interossei there are, Ist, a normal first palmar from the 2nd metacar- 
pal to the Index ; 2nd, a thin superficial palmar rising from the front of the 
8rd and 4th metacarpals, and inserted into the ulnar side of the first phalanx 
of the Index ; this muscle looks like an Adductor pollicis with a displaced inser- 
tion ; the other two palmar Interossei are normal. To the metacarpal of the 
pollex two small muscles are attached, one to the ulnar, one to the radial 
Bide, both springing from the second row of the carpus ; the inner of these may 
be an Interosseus primus volaris, or a flexor; the outer is evidently the 
adductor." 

' According to Duvemoy they are three in number (t. «. he reckons fonr, in* 
duding the Addnctor pollicis), and all ** adduct ** to the axis of the middle 
finger; they rise from the metacarpal bone of the digit into which they are 
inserted, in its whole length, and partly from the adjacent side of the adjacent 
metacarpal. It is plain that he only considers those as palmar Interossei which 
are inserted into digits which have no other slip of insertion from the dorsal 
Interossei on the same side, viz. the Index, ring, and little fingers. As, how- 
ever, several other slips, placed on the palmar aspect of the dorsal Interossei, 
and concealed by them from the dorsal aspect, are provided with distinct 



>.- 



MUSCLES AND NERVES OF A CHIMPANZEE AND ANUBIS. 189 

from tbe base of the adjacent one, and was inserted partly into the 
base of tbe tirst phalanx, partly into tbe extensor tendon, more dis- 
tally than tbe insertion of the dorsal Interossei. No. 1, chiefly from 
met. IL, partly from met. iii., inserted into the Index which it ad- 
ducted. No. 2, principally from met. iii., partly from met. ii., 
inserted into the middle finger, which it abducted radiad. No. 3 
rose principally from met. iii., partly from met. iv., and from an in- 
termuscular septum, described below, inserted into the middle finger, 
which it abducted ulnad. No. 4, principally from met. iv., partly 
from met. iii., and from an intermuscular septum at the origin of the 
Adductor pollicis, inserted into the ring finger, which it adducted. 
No. 5, principally met. iv., partly from met. v., and from an inter- 
muscular septum, described below, inserted into the ring finger, 
which it abducted. No. 6, principally from met v., partly from 
met. lY., and from an intermuscular septum, inserted into the little 
finger, which it abducted \ 

Dorsal irUerossei in An. (which strictly ought not to be called 
dorsal, as none had a dorsal origin) rose from the distal part of the 
palmar ligament, and also from the bases of the metacarpal bones 
into which they were inserted; the Ist ("Abductor indicis") having 
also an origin from the base of the metacarpal of the thumb. This 
muscle had also a raphe, which united it to the other indical (2nd) 
dor^•al Interosseus. They were 7 in number (as Church found the 
palmar set in the Cebus and Inuus nemestrinus); all the fingers had 
two, inserted on each side, and alternately abducting and adducting, 
except the little finger, which had one only. 

Palma/r rose from the distal part of the deep palmar ligament 
in common with the dorsal set They had a common origin, were 
flat, and concealed the subjacent ones from view. They adducted 
the little and ring fingers (inwards). According to Church's de< 
scription' of the Cebus and Inuus nemestrinus they resembled the 
same muscles in those monkeys generally, but differed in number; all 
were much like those of the Camivora. 

Some general remarks on the Interossei will be found under the 
description of those in the foot. 

tendons separately iDserted, it is better, I think, to desoribe the palmar Inter- 
ossei as six in number. Church found seTen in the Orang. 

^ The intermuscular septa mentioned above, from which Nos. 8 to 6 partly 
rose, were those between adjacent palmar Interossei. Since the palmar Inter- 
ossei in man are all adductors, sc. of the Index, ring, and little fingers, these 
were represented by the Ist, 4th, and 6th in the Chiup.; the abductors, so. 
two of the middl -i and one of the ring finger, were palmarly divaricated portions 
of the 2nd, 3rd, and 4th dorsal Interossei of man. They were, however, in my 
Ghimp. quite separate and palmarly placed. RoUeston describes them thus 
in the Omacp.: ** The dorsal Interossei have their two heads from the opposed 
faces of the metacarpals less intimately connected than in man, and the head 
from the metacarpal of the digit into which this muscle is inserted, sends its 
tendon into the fibrous extensor expansion on the dorsum of the digit, whilst 
the other head has its insertion into the basal phalanx, and having its origin 
placed dorsally, and connected with both metacarpals at their carpal end, has 
its insertion pidmarly to that of the other head.*' 

« P. 9. 



190 MB CHAMPNXTS. 

Did^hfragm was oarefiilly examined in Chimp., and was found 
strikingly anthi*opomorphic ; the 4 weak places (2 on each side), in 
which io man diapliragmatic or phrenic hernia sometimes occurs, 
viz. on each side of the ensiform cartilage, and also between the 
origin of the greater muscle from the last rib, and that from the 
Ligamentum ai^cnatum externum, were as well marked in the Chimp. 
as in man. 

Lower Limb. 

Femwr possessed a Ligamentum teres, in both Chimp, and An., 
as in all Quadrumana except the Orang. 

Faoas parvus in Chimp, rose from the last (13th) dorsal and first 
lumbar vertebrae, was attached to the bodies of the succeeding ver- 
tebrae bj a fascia which overlaid the Psoas magnus, and received from 
it and transmitted to Ps. p. a few fibres. It overlaid the Ps. m., and 
ended in a fiat tendon which was inserted into the lower part of the 
Ilium at the ilio-pectineal line, just where the femoral vessels emerged. 
In An. it was well developed, rose fix>m the intervertebral space 
between the Ist and 2nd lumbar, and from the £EU9cia iliaca and 
transversalis as far down as the 6th lumbar, where its tendon, which 
was very strong, began. It was continuous with Poupart*s ligament, 
and was inserted into the whole length of the upper border of the 
Pubes and lowest part of the Ilium, crossing the Psoas magnus as it 
left the pelvis. In man this muscle is very variable and inconstant, 
but geneiully rises from the last dorsal and first lumbar vertebrss. 

F8(Hu magnus in Chimp, rose h-om the last dorsal v., from the 
inferior surface of the interior inch of the last rib ; from the body and 
transverse process of all the (4) lumbar v. and the tendinous arches 
passing across the hollow formed by the constricted portions of their 
bodies, becoming continuous with the Iliacus below this origin. It 
was inserted with the Iliacus as in man. In An. it differed from 
that in man in rising not higher than the intervertebral space 
between the 1st and 2nd lumbar v. instead of from the last dorsal v. 
It was in two distinct parts ; an external and superior, rising from 
the intervertebral spaces between the 1st and 2nd lumbar v. and 
down as &ir as between the 5th and 6th lumbar v. supplied by the 
Anterior crural nerve ; and an internal and inferior, rising from the 
intervertebral space between the 3rd and 4th lumbar v. and as &t 
down as the 7th lumbar v. (this increased number of lumbar vertebrse 
accounts for many difierences both of muscles and nerves), supplied by 
the 3rd lumbar nerve. In man the whole muscle is supplied by the 
2nd lumbar nerve. The two divisions were separated by the course 
of the lumbar nerves. This furmshes another instance of reduplica- 
tion, and this set of muscles (the two Psoas and Iliacus) was much 
split up in this animal. 

Iliacus rose in Chimp, as in man, and soon fused with the Psoas 
magnus, the fibres running obUquely on each side into the Psoas 
magnus and its tendon, and gradually enveloping it from each side. In 
An. it was on the whole the same. Near it were two remarkable slips; 



MUSCLES AND NERVES OF A CHU^ANZEE AND ANUBIS. 191 

one, wbich was probably tbe same as that mentioDed by Owen' in 
the Orang (where, however, he does not mention any second origin 
from the Psoas magnos) was present on the right side, but not on the 
left in my young An., and altogether wanting in an adult An. which 
I examined speciaUy. It rose by two heads, one from the external 
factor of the Psoas magnus opposite the 5th lumbar v., the other from 
the exterior border of the Iliacus, just as it left the pelvic cavity. 
These formed a round tendon which was attached to the lesser 
Trochanter on the outer border of the rest of the Ilio-psoas. It 
resembled those described by Henle and Luschka as *' Iliacus intemus 
minor" except in having a psoas origin. It thus formed a Second 
lUo-psoaSy and so far justified Henle in using the term Ilio-psoas in 
man instead of Iliacus and Psoas. It offei's another interesting 
instance of reduplication. The second muscle rose from the Ilium 
at the under surface of the origin of the tendon of the Rectus, and 
from a line drawn from thence across the capsular ligament of the 
Femur. It was inserted into the line which runs from the. upper 
part of the linea aspeiu, spirally inwards and forwards, limiting the 
neck of the Femur, between the insertions of the Pectineus and 
Ilio-psoasw It was better developed on the left than on the right 
side. On the right side it was inserted beneath the " second IHo* 
psoas," and at the same spot (that musde being absent, as above 
stated, on the left side). It was a reduplication of the Iliacus, 
and, together with the iliac head of the "second Ilio-psoas," 
represents, or rather is represented, by the "Iliacus internui 
minor" of Henle and Luschka in man. The Ilio-psoad set of muscles 
was thus much split up (as were the Glutasi in the Chimp.); the 
Psoas magnus consisting of two distinct muscles, and there being 
a second llio-psoas, and a second Iliacus. 

Teruor vaginm /emoriB in Chimp, differed from that in man in 
extending farUier down the exterior border of the Ilium at its origin, 
in relation with the greater length of the Ilium. Its lowest point 
of origin was marked however as in man by the origin of the Sar- 
torius. It only reached the anterior superior spinous process of the 
Ilium after fusii%, or rising in common with the glutaeus medius and 
minimus externally. Its origin was also common to the Sartoriua 
internally and below, and the Transversulis internally and a,bove. In 
Ak. it rose half way down the anterior edge of the Ilium. 

OhiUcBua iryaai'fMw in Chimp, was smaller relatively than in 
man, but larger than in An., in correspondence with the more erect 
gait. It rose frx)m a fascia covering and giving partial origin to the 
Gluteus medius, and rising from the exterior part of the whole of the 
crest of the Ilium, and posteriorly continuous with the ftiscia covering 
the dorsal muscles; from the Sacrum, sacrosciatic ligament, Coccyx, 
and ischial tuberosity by an origin shared by the long head of the 
Biceps. (This ischial origin is described by Duvemoy as a separate 
muscle which he calls " Ischii-fSmorien.") Though it had this 
extensive origin, its muscular fibres did not reach higher than the 

1 Ftoc. Zool. S(K. Put I. (1880—81) p. 69; see also Ghnreh, p. 16l 



192 MR CHAMPNEYS. 

sacro-sciatic notch ; above this point what is mu-scnlar fibre in man 
wem fibrous tissue in the Chimp. It was separate from the Tensor 
vaginie femoris. It was inserted into the whole length of the linea 
aspera along the origin of the Vastus eztemus (Henle mentions that 
in man some of the most external fibres are inserted alongside of the 
Vastus extemus), and also by a distinct tendon into the fiiscia lata 
just below the great Trochanter. Its texture was coarse, especially in 
its lower part. Part of its tendon gave origin to some fibres of 
the Vastus extemus and short head of the Biceps, which in man rise 
from an intermuscular septum occupying the same position, and which 
I think serve to identify that septum as the homologue of the tendon 
of the GlatseuB maximus of the higher monkeys. A similar instance 
I have recorded under the Dorso-epitrochlien and Pronator radii 
teres. In An. it was covered by a dense fiiscia containing much 
hard yellow fat and clinging very close to the muscle. It rose not 
higher than the lowest part of the Sacrum and the root of the tail. 
Its fibres became fi^sed with those of the Tensor vaginss femoris 
opposite the great Trochanter (at which point some fibres were given 
off to terminate in the linea aspera), while the remainder terminated 
with those of the Tensor vaginae femoris in the fascia lata. The 
muscle was weak, especially at its origin. 

OlvJtceua medius in Chimp, was the largest of the three Gluteei, and 
relatively larger than in man. It rose from the fossa Ilii below the 
crest in its whole length, and as far d<jwn as half the length of the 
Ilium. Also, opposite the upper fourth of the Ilium, from the 
fiiscia above-mentioned, overlying it. It was inserted into the poste- 
rior edge of the great Trochanter in its whole kiigth. A small 
&sciole a, quarter ojf an inch broad separated from its anterior or exte- 
rior border, and was inserted into the anterior and distal part of the 
great Trochanter, on the opposite side of the Vastus externus from. 
rest of the glut. med. In An. it was largely developed as usual in 
Quadrumana, and with difiiculty separable from the Pyriformis. 

GhUcnia minimtiSj the smallest of the three in the Chimp. 
(Macalister found it twice as large as the Gluteus medius), rose from 
a line drawn from one inch below the ant. sup. spincfus of the Ilium, 
to its posterior and inferior extremity, opposite the acetabulum. The 
fibres from the anterior or superior portion rose from a wider origin 
than the rest, the line of origin decreasing in breadth as it ran back* 
wards. It had no origin from the coccyx, it could be separated 
with some difficulty into two nearly equal portions, an interior and 
deeper, and a posterior and superficial, overlapping the hinder part of 
the first. From the former of these the Scansorius was differentiated. 
The tendons, which occupied equal lengths of insertion along the 
proximal or upper 1^ in. of the anterior surface of the great Tro- 
chanter, were distinguishable but not separate. These two divisions 
are mentioned in man by Henle. Beneath the Glutseus minimus & 
small muscle a quarter of an inch broad, and two inches long, quite 
separate from it, rose from the exterior edge of the Ilium on a level 
with the uppermost part of the ischiadic attachment of the sacn>- 
sciatic ligament, and was inserted into the anterior edge of the great 



MUSCLES AND NEHVEd OF A CHIMPANZEE AND ANTJBIS. 193 

Tit)chanter just opposite the insertion of the second part of the 
Glutaeus medins with which it agreed in breadth. In Ak. it showed 
no signs of fission or reduplication; it rose from the exterior of the 
Ilium within two lines, the upper starting from a point one-third 
down the anteiior edge of the Ilium, and running backwards and 
downwards to the upper extremity of the sacro-sciatic notch; the 
lower running from a point two-thirds of the distance down the 
anterior edge of the Ilium backwards and downwards to a point just 
opposite the acetabulum. It was inserted as in man. 

The Scansorius in Chimp, was very imperfectly separated from 
the Glutffius minimus, the anterior half of which overlapped it. It 
formed about a quarter of that muscle, being differentiated from the 
anterior and deep portion, and was inserted together with its most 
anterior poi-tion into the anterior and distal part of the great Tro- 
chanter. It was supplied by the superior Gluteal nerve in common 
with the rest of the Glutseus minimus ^ In An. it was undifferen- 
tiated'. 

Pyriformxa in Chimp, was really, but not obviously, separate from 
the GlutflRus medius. It rose from the lower part of the 2ndy 
3rd, and 4th, ^nd upper part of the 5th sacral vertebrse, from the 
adjacent part of the deep surface of the Ilium, but not from the sacro- 
sciatic ligament. It was inserted into the proximal end of the great 
Trochanter, being at its extremity slightly united with the tendon of 
the Glut med., with which it shared a bursa mucosa. In An. it was 
nearly fused with the Glut. med. 

Ohtv/ralor interrvas in both had an origin somewhat more extended, 
and elongated by the increased length of the iliac bones than in 
man ; viz. frt)m the whole of that part of the brim of the pelvis 
which was formed by the Ilium and Pubis, except opposite the obtu- 
rator foramen, where the fibres rose from the tendinous arch which 
ran below the obturator vessels and nerve, across and below that 
foramen ; from the internal a8i)ect of the long symphysis Pubis, and 
adjacent part of the lower border of the Ischium as far as the tu- 
berosity ; also from the obturator membrane, and from all the inter- 
nal surface of the Pubis and Ischium mesiad of a line drawn perpen- 
dicularly through the obturator foramen. Its tendon was overlapped 
by the Gemelli and fused with them, and they were inserted together 
into the digital fossa on the internal and posterior side of the greab 
Trochanter, as in man. 

Gemelltis superior in both rose from a point just above the troch- 

' Yrolik could not find it. Kacalister found it one-fifth of the size of the 
Glatffius medins. Its presence in man is recorded by Wood. 

* In Chimp, we see a remarkable tendency in the Gluteal set of muFoles 
towards fission or reduplication, such as we found to obtain in many other 
mnsoles, which we have noticed as they have been described, but in An. 
especially in the Ilio-psoad set : — the Gluteus maximus in Chimp, was so far 
separated into two as to have been described by Duvemoy as two muscles ; the 
Gluteus medius was bifid and had its two parts differently inserted; the 
Glutffius minimus not only showed the two diyisions described in man by Henle, 
but in addition threw off a Scansorius, and a still more distinct muscle men- 
tioned above, from its deep surface, thus being split into four divisions. 

VOL. VI. 13 



194 MB CHAMPNEYa 

lear snrfiuse of the IsGhlam, where there was a Bmall mdiment of 
the ischial spine. 

GemeUua inferior in both as usually in man, much the smaller 
<^ the two GremellL It rose from an origin 2 in. in length, conb- 
mencing at the most dorsal, posterior, or superior part of the inter- 
nal ridge of the ischial tuberosity, and ending at the rentral, inferior, 
or mesial one-thii*d of the same edge. 

QtutdrcUtis /emoris in both arose as in mau. It was inserted 
not into the linea quadiati as in man, but into a horizontal line, 
extending from the lesser Trochanter outwards, and a little down- 
wards for about three-fourths of an inch. There was, however, about 
half way along this line a small yertical insertion extending along the 
posterior edge of the great Trochanter, in a line feebly represent- 
ing the linea quadrati of man. The upper pai-t of the insertion of 
the Adductor brevis OTorlapped the exterior part of the insertion of 
this muscle. 

Coccygeus in Cmicp. was as in man, except that it was readily 
divisible into two portions. Its insertion was fused with the coccygeal 
part of the origin of the GlutsQus maximus. The perineal muscles 
had been destroyed in removing the abdominal viscera^ 

Biceps femoria in Chimp, was as in man in general. The short 
head was well developed; the long head rose, as in man, from the 
most external part of the external edge of the tuber Ischii, its origin 
being common also to the Semitendinosus, but also, unlike man, 
to the Semimembranosus which the Semitendinosus overlapped; to 
the most posterior part of the Gracilis, which the SemimembiaDoeus 
overlapped and which was much larger relatively than in man, also 
to the lowest part of the origin of the Glut, max., which in man has 
no origin £rom the tuber IschiL Its tendon ran for one inch be- 
fore receiving the fibres of the short head. The fusion between the 
tendons of the two heads was not complete, but the tendon of the 
long head crossed over that of the short head and was inserted into 
€he out-er and anterior tuberosities of the Tibia, as well as into the 
fascia of the 1^ which was continuous at the knee-joint with the 
fascia lata of the thigh. The tendon of the short head crossed under 
that of the long h^id and was inserted into the prominence at the 
exterior side of the head of the Fibula, and into the fascia of the leg, 
distally to the tendon of the long head\ In An. the Biceps was very 

^ Thd proper tendon of each of the heads was \ in. broad. In the Qorilla and 
in the Orang the Bleeps consists of two distinct mufides, no fusion taking place 
as in Chimp, and still more in man. In Ohimp. compared with most other 
Qnadrnmana the insertion was very high, and the masde itself small, an 
antibropomorphic point. In man there is but one conjoined tendon from the 
two heads. The original separateness of the two heads was even in Chimp. 
plainly indicated by the difference in the mode of their innervation ; they were 
both supplied by the great Sciatic nerve as in man bat in a different way. In 
man this nerve gives off separate branches to the Adductor magnus, Semi- 
tendinosus, Semimembranosus, and Biceps. In Chimp, a separate trunk was 
formed as the great Sciatic issued from the sacro-soiatio foramen, and after a 
course of 4 in. gave off (1) a branch which supplied the origin of the Semitend. 
by several twigs, (2) a branch which bifurcated and supplied the upper one-third 



tftJSCLES XSJ> NfiKYES OF A CHIMPAKZEE AND AKUBIS. 195 

large. Its short head was wanting, as sometimes in man (Henle and 
Theile) and the lower monkeys. Its insertion was very long, occupying 
nearly half of the Fibula. It was inserted chiefly into the fascia of 
the leg, a strong band being given off to the covering the knee-joint, 
and a less strongly marked one to the outer tuberosity of the Tibia, 
but there ap|)eared to be no special insertion into the head of the 
Fibula \ 

Semitendinosvs in Chihp. rose from the tuber Ischii in common with 
and below the long head of the Biceps as in man, with which it was 
fused for its first three inches, and also in common with and superficially 
to the Semimembranosus, unlike that in man. It differed fi-om that 
in man in being larger instead of smaller than the long head of the 
Biceps, and not having a tendinous inscription (which, however, 
was found by Macalister in his Chihp.), also in its insertion, which 
was comparatively much lower down than in man. Yrolik found it 
inserted as in man. Its tendon proper was inserted into the anterior 
tubercle of the Tibia, two inches from the top of that bone, and was 
well marked, flat (it is round in man), and one-third of an inch broad; 
but about three inches before it reached its insertion it suddenly gave 
off a wide-spreading expansion (represented in man as observed by Ellis, 
p. 705) downwards, which was three-fourths of an inch wide at one 
inch distant from its origin, and the most posterior fibres of which 
became quite perpendicular. This expansion became continuous with 
the fascia of the leg. The insertion was overlapped by that of the 
Gracilis, with the aponeurotic expansion of which it fused. The 
tendon was not so long as in man, as Yrolik also found*. In As, 
this muscle rose in common with the long head of the Biceps, and 
was inserted very low down the 1^, much lower than in Chimp. 
Two principal tendons were given off, one exactly opposite the upper 

of the long head of the Biceps, (8) it split into branohes which supplied the 
proximal part of the Add. ma., tl^e mesial part of the Semitend. and Semi- 
membr. and the separate external or distal divisiou of the Add. ma. The short 
head was supplied by two twigs separately rising from the main trunk of the 
great Sciatic nearly opposite the middle of the thigh. The bearing of this 
arrangement on the general question of progress as indicated by Integration 
will be found noticed under the great Sciatic nerre. 

^ The distinctness throughout of the two heads and their tendons in the 
Gorilla and Orang, as well as their different mode of innervation in Chimp. 
and the absence of the short head in the Oebus and Inuus nemestrinns, in 
An. and many other Quadrumana, and also occasionally in man, all point to 
the essential distinctness of the two as separate muscles. The progress from 
the absence of the short head in the Cebus, Inuus nemestrinus, An, &c. , and 
the complete distinctness of the heads and their tendons in the Gorilla and 
Orang, through the partial fusion of the tendons in Ohihp., to the complete 
fusion in man, is remarkable, and would, as far as it goes, serve as an argument 
lor placing Chimp, at the head of the Quadrumana. 

* Cuvier has remarked that in all Mammals below man this muscle and the 
Semimembranosus possess this aponeurotic expansion at their insertion, and 
that their insertion is also muoJi lower down the leg than in man, which 
keeps their knee necessarily bent and is incompatible with an erect gait. The 
approach to an erect gait is therefore indicated by the removal upwards of their 
insertions. Professor BoUeston informs me that this comparatively low inser- 
tion of the hamstring muscles is still to be seen in yoimg children — a most 
significant fact. 

13—2 



196 MB CHAMPNEYS. 

part of the insertion of the Gracilis, and inserted into the lower part 
of the anterior tubercle of the Tibia^ the lower fusing with the lower 
end of the Gracilis, and ending with it in the &scia of the leg at a 
point more than half way down the leg. There was no teudinous 
intersection. 

Semim&Tibranosus in Cbimp. rose as in man. It was overlaid 
at its origin by the conjoined origin of the Biceps and Semitendioosus, 
^and was fused with that part of this conjoined origin which was 
continued into the Semitendinosus for one inch, and after that, partly 
(by several small and separate tendinous slips) for another 1^ inch. 
Its tendon of origin was flat and long, as in man, being 3 inches 
in length. The muscle was of the same size as the long head of the 
Biceps, i,e, smaller than the Semitendinosus; in man it is larger than 
either. Its insertion differs from that in man (Macalister, in p. 349), 
in not possessing a slip expanding into the aponeurosis overlying the 
popliteus muscle. Between its tendon and the internal lateral liga- 
ment of the knee-joint was a bursa. It sent no fibres to this ligament 
as it does in man. Its tendon of insertion was rounded and small, and 
reached the Tibia one inch more proximally than the upper part of 
the insertion of any of the other three hamstring muscles** In An. 
it resembled that in man, except that it had no membranous origin 
and only one insertion, viz. that into the posterior part of the inter- 
nal tuberosity of the libia. 

GracUia in Chimp, rose from the whole length of the Symphysis 
Pubis, and the interior inch of the upper edge of the pubic ramus, 
by a flat membranous tendon half an inch long, which overlapped 
diagonally and fused with the part of the Add. 1. which was adjacent to 
the most exterior part of its origin. It was inserted just superficially 
to the insertion of the Semitendinosus and agreed with the latter in J^ 

the breadth of its tendon proper (half-inch), but differed in having its 
tendon a little the shorter ; it also agreed with the Semitendinosus 
exactly, in the insertion of its tendon proper into the lower part of 
the anterior tibial tubercle, 2 in. from the top of that bone, and of a 
itipidly spreading tendinous expansion into the fascia of the leg. It 
diffei*ed quantitatively in an enormous degree from that in man in 
every particular, being much larger, as is the case in the orang 
(Church, p. 10), but agreed qualitatively. Its insertion, like that of 
all the hamstring muscles except the Semimembranosus, was much 
lower than in man. It was the largest of ikie hamstring muscles^ and 
was broad and flat. In An. it had a wide origin and was not inserted 
into the inner tuberosity of the tibia, but into the fascia of the leg 
and anterior ridge of the middle third of the Tibia, straightening 
of the limb. 

Sartorma in Chimp, was large, and rose from the lower part of the 
anterior edge of the Ilium, and was fused with a few of the fibres of 
the external part of the Ilia'^us, with Poupart's ligament, with the 
Tensor vaginse femoris, the Gl. min. and the Kectus femoris', inserted 

^ Vrolik says it is inserted lower down than in man, but this was not the 
case in this specimen, nor did Owen find it so. 

* No such slip from the origin of the Pectinens, passing nnder the femoral 



MUSCLES AND NERVES OF A CHIMPANZEE AND ANUBIS. 197 

Buperficiallj to the Gracilis, but without a bursa between them. In 
Ak. it rose as in man, but was inserted into the upper half of the 
anterior ridge of the Tibia and the fascia of the leg. 

Rectvs femoria in Chimp, resembled that in man, but rose by one 
and not two separate tendons as in man \ inserted as in man, but was 
more separate from the Crureus than in man. In An. there were 
also not two separate heads of origin ; the fibres had no penniform 
arrangement. 

Vastus extemvs in Chimp, agreed with that in man in every 
respect, except in its relations to Glut max. (which see above). 

In An. it was, as in man, except that the short head of the Biceps 
being absent, it bad no origin from an intermuscular septum between 
the short head of the Biceps and itself. 

Vastvs irUemits in Chimp, rose higher, as far up as the neck of 
the femur. 

Crureus in Chimp, differed in being more closely connected with 
the Vastus ext. than with the Vastus int. 

Subcrureus in both was absent. 

Pectineus in both as in man. 

It was not bUaminar in either, but it was found by Macalister to 
be so in Chimp. 

Adductor longus in Chimp, was generally as in man. In An. it 
arose as in man, but differed slightly in falsing with the tendon of 
the Adductor magnus. 

Adductor brevis in both, was in general as in man but rela- 
tively larger. It was mtlllti-fasciculate, but in Chimp, was inserted 
into the oblique line leading not from the small but the great Tro- 
chanter, to the linea aspera. The Obturator nerve pierced it and 
divided it into two portions, at the same time supplying it as in man. 

Adductor magnvs in Chimp, consisted of two distinct divisions, 
the deeper and larger rose from the whole of the anterior surface of 
the pubic bone along the Symphysis, and the adjoining part of the 
Ischium as feir as the tuberosity. It was inserted into the linea 
aspera along the lower half of the thigh, fusing with that of the 
Add. 1. It was multi-fasciculate, and supplied by the Obturator nerve. 
The superficial and smaller division, more compact, overlapped the 
preceding below, and arose from the Tuber Ischii by a flat thin ten- 

yessels, as is described by Owen, conld be found, but with regard to this con- 
nection when it exists, Guvier*s remark on the ** castor *' (beaver) is not a little 
interesting: "Le couturier est tout-li-fait confondu dans le castor avec les 
pectin^s." 

^ Yrolik asserts that there are two origins ; — in my Chimp, the tendon of 
origin arched round the upper edge of the acetabulum (as does the 2nd origin in 
man), but this portion was not separate. In man, however, the two origins 
are united by membrane, and in Chimp, they admitted of being readily sepa- 
rated. It rose from the anterior inferior spinous process of the Ilium (as m man) 
and from two lines diverging from it on each side like the legs of the letter A, 
and capping the acetabulum, also from an aponeurosis giving origin in order 
from above downwards to the Tensor vaginiB femoris, Glutaeus minimus, Sar- 
torius and Bectus, and, externally to the origin of the Bectus, to the small 
muscle mentioned as underlying the GlutsBUs minimus ; by which aponeurosis 
the origins of all the above-mentioned muscles were ooxlnected. 



198 .MB CHAMPNETS. 

don, and was inserted separately into quarter of an inch on the internal 
side of the internal condyle of the Femnr\ In An. it was multi- 
fasciculate, but not in two separate divisions. 

Obiuralor extemus in Chimp, had its origin, in genera], as in man. 

Gastrocnemvua in Chimp, was as in man and was lai^e, but the 
muscular fibres were continued as far as the insertion into the calca- 
neum ; see altK) foot-note'. 

SolcBU8 was large. It rose from the head and upper three-quarters 
of an inch of the Fibula instead of from its upper one-third. There 
was no origin from the Tibia'. 

Flantaris in Chimp, as in man. It was absent in the right leg\ 

In Aif. it was fused with the outer head of the Gastrocnemius for 
half an inch from its origin, and was partly united with it by ten- 

^ Gnvier in his plates draws in a figure of a Magot, an Adductor having a 
sixnilar but not so distinct insertion (I'), which he cidls **long addncteur " but 
as he also calls it '*l8chii-f^morien" it cannot be homologous with the Add. 1. of 
man (Church has, wron^y I think, translated it "Adductor longus," p. 18), 
which rises from the Pubis. This fascicle is apparently the same as that here 
described in Chimp. Ellis describes two more or less distinct parts of this 
muscle in man, which differ in the same way in texture and insertion, but are, 
not really separate. In Chimp, the superficial part of the Femoral artery 
divided the two portions before reaching the popliteal space; and since it 
pierces the Adductor magnus at the same portion of its course in man it furnishes 
another reason why the distinct second portion in Chimp, may be identified 
as part of the Add. magnus, specialised. Moreover, the Obturator nerve sup- 
plied the principal portion, the great Sciatic nerve the superficial and smaller 
division, both of these nerves in man supplying the Adductor magnus. Henle 
describes a slip somewhat similar to the second portion which I have described. 

Burdach describes the 3 Adductors as one muscle in 5 divisions ; the Add. 
m. furnishing two, the Add. 1. the 3rd, and Add. br. the 4th and 6th. He also ^ 

says that Meckel includes the Pectineus as a 6th (could he do so consistently if # 

he had found it bilaminar 7). 

' Church says that in the Orang it is small and often separate from Solsus; 
in Inuus nemestrinus it was more separate than in the cases in which it was 
found to be fused in the Orang. The only points in the Chimp, worth remark are 
that the inner head was fused with the insertion of the second portion of the Add. , 
this head neither being separate nor extending so low as in man. A bursa 
underlaid this head and communicated with tide knee-joint as in man. No 
sesamoid bone or fibro-cartilage could be found in the external head. The 
tendon slightly differed from that in man, in that tiie muscular fibres were 
continued as far as the insertion into the Calcaneum (Macalister's Chimp. 
differed in this respect from mine, while Wilder*s agreed with it), running on 
each side into the tendon whi<di lay in the middle. 

* The tibial head was absent in a Chimp, dissected by Macalister (the 
fibular origin being very large). It ia absent in An.; in the Orang and 
Cebus (Church, p. 14), and the Gorilla (Duvemoy, p. 93). In the Inuus 
nemestrinus it rose from the fibula and external condyle of the Femur. Yrolik 
mentions in Chimp, a tibial but no fibula origin. The tibial head was 
found by Humphry in a Chimp. The fibular origin was found a mere slip in 
Chimp, by Humphiy and Huxley, and by Church in the Orang. 

* Macalister found it very small, present in ike left leg, absent in the right, 
the opposite arrangement was found by Wilder. Vrolik found it as well as 
Huxley and Humphry; it was absent in Traill's specimen. In the Orang 
(Church, p. 14) it was absent, also in tiie Cebus ; it was large in the Inuus 
nemestrinus and arose as in the Chimp. Duvemoy says it is absent in the 
Gorilla and Orang. It is often absent in man. It was absent in a young Chimp. 
dissected by Dr Embleton. 



MUSCLES AND NERVES OP A CHIMPANZEE AND ANTJBIS. 199 

• 

dinons dips for one inch further, at this point being connected with 
the outer head of the Gastrocnemius by a broad fibrous hand. Its 
muscular belly was longer than either of those of the Gastrocnemius, 
and its greatest breadth was half that of the SoUbus'. 

FoplUeus in Chimp, as in man. 

In An. a small slip of muscle, not found in an adult An. specially 
examined, but present in a Wanderoo (Simia ferox), ran firom the 
upper (external) head of the Popliteus to the internal head of the 
Gastrocnemius. 

FUaooT longiu digitorwn in Chimp, arose as in man. Its tendon 
was more fused with that of the Fl. 1. h. It could be seen, that, but 
for the fusion above mentioned, this muscle would fumi^ tendons' 
only to the 2nd (index) and 5th toes'. 

In Ax. it was the sole mover of the index and little toes, but 
moved the others by its intimate connection with the Flexor longus 
hallucis. It also sent a distinct slip to the tendon of the Flexor longus 
hallucis going to the hallux. 

Lwmbriciiea. Most of these muscles in Chimp, took origin both 
from the tendons proper of the FL 1. d., and those of the Fl. L h. That 
one, however, which went to the 2nd toe rose only from that of the 
FL L d. It was the largest. That one which went to the 3rd toe 
rose by two heads, one fit>m the tendon of the Fl. I. d. going to the 
2nd, principally from the fibular, but partly also from the deep and 
tibial aspects ; the other from the tendon of the Fl. 1. h. going to the 
3rd toe, from the internal, and internal half of the superficial surface. 
That one which went to the 4th toe rose from the tendons of the 
FL L h. going to the 3rd and 4th toes, from the adjacent halves of 
their superficial aspects, and the adjacent sides. That one which went 
to the 5th toe rose from the tendon of the Fl. L h. going to the 4th 
toe, from its superficial aspect, and fibular side, and by another small 
belly, from the adjacent half of the superficial surface and tibial 
side of the tendon of the Fl. L d. going to the 5th toe. Most of 
these reached proximally as far aa the division into separate tendons, 

^ In oQntraBting the mnsdles of the tendo AohiUis in OmMP. and An., 
we find the general arrangement very anthropomorphio in the former, very mnch 
the reverse in the latter, hnt there are seme exceptions. For while Axr. 
has a smaller Gastroenemius, the heads of which are separate far down, on the 
other hand its tendon is longer; the mosoolar fibres quite ceasing half way 
down the leg, though An. was a young one. In both the SoLbos has only 
a fibular origin. The Plantaris is large and partly fused with both the Solnus 
and both heads of the Oastrocnemios in An., but is small and separate 
except at its origin in Chimp, as in man. In An., however, the tendons of 
all three mnades remain separate till just before their insertion, when they 
simnltaneoasly fuse; in Chimp, as in man the Solsns is fused during 
nearly its whole extent with the Gastrocnemius, the Plantaris being separate at 
least in its muscular portion; its tendon also not fusing completely till just 
before insertion. 

' Duvemoy describes in the Gorilla the Flexor longus digitonun sending 
tendons to all the digits. Church found it in the Orang sending tendons to the 
2nd, 4th and little toes, and sending no slip to the tendon of the Flexor longus 
hallucis. In my Chimp, it was inserted as in man, and in the same way as the 
Flexor profimdus in the hand. 



200 * MR CHAMPNEYS. 

and extended distally for three quarters of an inch, but the small 
belly of the Lumbricalis of the little toe had only a minute origin 
which was attached about three quarters of an inch from the division 
of the tendon of the Fl, 1. d. going to the little toe. 

They were inserted as in man, and were quite rs well developed 
as those of the hand, in which point they differed from those in man, 
which in general they resembled. The chief differences were those of 
origin, which were due to the continuance of tendons from the 
Fl. 1. h. to all the digits except the index and little toes^ 

Flexor cuxesaorius in Chimp, in both feet rose from the Cal- 
caneum^ a little anterior to the internal tubercle (by one head, not 
two as in man) and stretching inwards aud forwards was inserted 
into the external edge of the tendon of the Fl. 1. d. just before the 
fusion with the tendon of the Fl. 1. h. The teudon was much louger, and 
smaller than in man ^ In An. it was well-developed, and was present 
also in an adult An. specially examined, and rose from the fibular side 
of the middle part of the plantar surface of the Calcaneum by a fleshy 
head, and from the adjacent corner of the Cuboid by a tendinous 
head; and was inserted into the outer side of the point of inter- 
communication of the common tendons of the Fl. 1. d. and 1. h., thus 
mnning diagonally across the Calcaneum. 

FUxor hrev^is digitorum in Chimp., a very complicated muscle. 
The principal portion rose from the inner side of the os-calcis as far as 
the tuberosity, and from the deep surface of the plantar fascia, by 
which it was connected with the origin of the Abductor pollicia. 
Two minute tendons were sent from that going to the third to 
fuse with the tendon of the flexor brevis going to the second (in- 
dex) toe, just mentioned, which they did opposite the metacarpo- 
phalangeal articulation, one of them developing about half-way a 
very small muscular belly. A small muscular belly was also detached 
from the main portion, and ended in a tendon which fused with the 
tendon to the 4th toe. This last rose from the surface of the 
tendon of theFL 1. d., as far as the internal malleolus, and was chiefly 

^ Duvemoy says, that only the Lumbricalis of the 2nd toe arises from the 
corresponding tendon of the Flex. long, dig., the others rising from the tendon 
of the Flex. long. hal. Dr Embleton mentions " a small muscle accessory to 
the LombricaleB arising from the long Flexor tendon before its division." He 
gives no further acconnt of it. Could it possibly be that part of the Fl. br. d. 
which sent a tendon to the 4th toe in mine ? 

' This muscle was absent in Bolleston's and Embleton's specimens. Hnm- 
phry found it small in both feet of one Chiicp. In another Chimp, it did not 
reach the flexor tendon in one foot, and was absent in the other foot. Church 
found it in the Orang, sending a tendon to the tendon of Flexor longos to the 
little toe, and another, which accompanied that tendon, and, after being per- 
forated by it, was inserted into the second phalanx of the little toe. Humphry 
could not find it in Orang, but found it large in Ateles. In An., Cebus, and 
Inuus nemestrinus, it fused with the tendon of the Flexor longus digitorum, as 
in man. 

The coexistence of this muscle with the irregolar slips described under Fl. 
br. d., which have been stated (as by Vrolik, and apparently by Church, as 
above) to partly represent the Accessorius, tends to establish their nature 
as that of scattered portions of FL br. d., as hereafter described, and at 
any rate in Chimp, disproves their homology with the Moles camea. 



MUSCLES AND NERVES OF A CHIMPANZEE AND ANUBIS. 201 

inserted into the 4th toa A portion of it joined the tendon 
bef«jre described going to the 3rd toe. A small slip also rose from 
the tendon of the Fl. 1. d. to the little toe, and was inserted iudis- 
.tinctlj, being mnch fused by fibrous tissue, with the tendon of the 
FL 1. d going to the little toe. The fascicle to the little toe is most 
irregular in its arrangement in the Quadrumana and in man. 

In An. the Fl. br. d. was very different from that in man. It 
consisted of the following parts : (1) a long head rising from the 
lower Bur&ce of the Calcaueum, iti common with the Abd. p. and 
Abd. 5ti as well as to fascicle 2. It was inserted into the 2nd toe. 
In the Ibft foot but not in the right, it received two minute slips 
from the next (2) A compound fascicle from the conjoined tendon of 
Fl. 1. h. and Fl. 1. d.,and consisting of one proximal and distinct belly, 
and three other distally placed and less distinct bellies; they ended in 
a tendon inserted into the 3rd toe. In the right foot, but not in the 
left^ the first and second bellies of poj-tion 2 each gave a small 
tendon to the conjoined tendon of Fl. 1. h. and Fl. 1. d. (3) from a 
similar and parallel origin inserted into the 4th toe. The last two 
received slips from the plantar fascia. (4) A small fascicle in the 
left foot, from a belly common also to portion 3 (in the right foot 
rising separately), inserted into the 5th toe. 

TibicUis posticue in Chim. rose as in man. Its tendon split 
more definitely than in man into two, one of which was inserted into 
the Scaphoid, the other into the Ecto-cimeiform bone. Ko sesamoid 
body oould be found in its tendon, as is usual in man. In An. as in 
man, except that its tibial origin did not extend so far down as that 
of the Fl. L d. 

Flexor Umgtis haUucia in Chimp, rose as in man, except that the 
origin of the Solseus not extending down the Fibula for more than 
one inch, all below this was occupied by it. Its tendon, besides fur- 
nishing a tendon to the Hallux, furnished one to the 3rd and 4th 
toes*. In An. this muscle rose equally from Tibia and Fibula, from 
the latter of which in man it is separated by the Tib. post. This 
tibial origin, which was paler, explains the fact that the Fl. 1. p. rises 
in man from the radius, the Fl. 1. h. from the Fibula; we here, 
as it were, see the muscle transferring its origin*. The tendon was 
intimately fused with that of the Fl. 1. d., so that the action of either 
muscle bent all the toes. As usual in Quadrumana it supplied the 
middle and fourth toes as well as the Hallux. The tendon to the 
Hallux, which passed through a ligamentous ring giving partial origin 
to the inner head of the Flexor brevis hallucis, received a slip from 
the tendon of the Fl. 1. d. Theile mentions a similar slip in man. 

Feroneus longus in Chimp, as in man, but strong, and the fleshy 
fibres extended to the malleolus. In An. it was as in man, except 
that it rose from the upper half of the Fibula, instead of from the 
upper third. 

^ The doBoriptions by varioufl authors shew that these two muscles in Quad- 
rmnana vazy greatly in their relations to one another and in the toes they 
respectively sapply. See Vol. i. of this Joum. p. 266. 

' In man this muscle is very invariable. Henle, p. 292. 



202 KB 

Peroneus hrev. in Chdcp. reaembled that in man in its ori|;in, 
Imt was stronger, and had a second tendon running along the 5th 
metatarsal connected bj fibrous tissue with that bone, and fiudng * 
with tendon of Ext. 1. d. and lumbricalis. This second insertion is 
not uncommon. In Ak. as in man, but rose fi\>m the middle l-3id 
instead of the lower half of the Fibula. 

Peroneus tertiua absent in Chimp.^ In As. it. differed consider* 
ably from that in man. On both sides it rose from nearly the middle 
one-third of the Fibula enclosed in the P. br. Ha tendon passed 
through the annular astragalo-calcaneal ligament with that of the P. 
br. (the Per. long^ being in a separate channel). Its tendon fused 
with the extensor tendon of the little toe. This will be seen to 
be really a Peroneus quintL 

Extensor longus digitorum in both as in man*. 

BocUneor proprius halhtcie in Chimp, as in man, but more power- 
fully adapted for abduction by passing under another ligament, besides 
the annular ligament, which extended frt>m the tuberosity of the Sca- 
phoid to the base of the inner metatarsal bones, and which gave 
passage also to the two tendons of the Tib. ant.* In Ak. it rose from 
the upper two-thirds of the Fibula and interosseous membrane, but 
otherwise as in man. 

Tibialis aaUicus in Chimp, was, ss is often the case, double^ 
and the internal and larger tendon inserted into the Ento-cuneiform 
bone was separate throughout from the external and smaller tendon, 
which was inserted into the base of the metatarsal bone of the 
Hallux. In Ak. it rose only from the upper half of the Tibia. It 
showed a tendency to become split, and we must remember that this 
was a young animal. 

^ SCftcalister says it is never present in Qnadramaiia, "the so-called Pero- 
neus tertius of Wyman, in the Howling monkey, being a Peroneos qainti f 
Bolleston, however, found it in a Chimp. Churoh mentions it in the Innus 
nemestrinns and Cebns, and I fonnd it in Ak. on both sides ; bat Iq all the 
three latter at least it did not pass together with the tendon of the Ex. 1. d., 
bat with that of the P. br. Again, it was not inserted into the base of the 
fifth Metatarsal, but fused with the tendon of the Ext. 1. d. oppoeite the 
Metatarso-phalangeal articulation, jast as did the accessory tendon which I found 
to the Peronens brevis (see above). In Ihe Cebus it perforated the tendon 
of the Peronens brevis opposite tne Cuboid bone. Does not this perfora- 
tion in the Cebus of the P. br. tendon, together with the second tendon 
which I found to the P^oneus brevis in Chimp., go to show that the bo- 
called P. tertius of Quadrumana is really a ^vaiication of the P. br. ? Wood 
mentions a " Peroneus quinti" as a human anomaly. 

' The tendon, after passing under the anterior annular ligament, passed 
through a separate sheath springing from the base of the Caloaneum, and 
again iuserted close to its origin, running upwards and inwards; from the 
superficial end of this sheath a small slip ran inwards to join the annular 
ligament over the internal malleolus. I can find no description of this liga- 
ment in any of the books, but I found it in An. and Ch. By its means Uie 
tendon was held just distally to the external malleolus. It is not found in man. 

' This ligament is mentioned by Duvemoy in the Gorilla, but he says that 
the tendon follows the line of the metatarsal and first phalanx of the thumb, 
which was not the case in my Chimp. It is not mentioned by Yrolik, Mao- 
alister, or Church, nor can I find any mention of it in Chimp. It was pre- 
sent in An. — Henle mentions that in man this musde is occasionally double. 



/ 



HUSCLES AND NERVES OF A CHIlfPANZEE AND ANUBIS. 203 

JSostensor hrevia digiiartbm in Chimp, was as in man, except that 
it rose partly from the special sheath from the Calcaneum, trans- 
mitting the tendons of the Extensor 1. d. The hallucal division 
was not separate, as has frequently been fonnd by others to be the 
case. In An. it was as in man, except that the tendon to the Hallux 
and next toe were given off by a common muscular belly, and the 
inner side of the muscular belly for the middle toe was attached by 
£iscia to the inner side of the metatarsal of that toe, besides its 
insertion into the tendon of the common Extensor. A small muscle 
rose from tbe Calcaneum at its neck, and was inserted into the inner 
side of the base of the metatarsal bone of the great toe, together 
with one of the tendons of the Tib. ant It exhibited a variability, 
which we found to obtain in the muscles of the hand, being absent 
in an adult An., specially examined. It would seem to be a redu- 
plication of the Extw br. d., such as we have noticed in other muscles, 
e.g. Abd. p., Fl. br. 5ti (in the hand), and liiacus and Psoas in the 
lower limb. 

Abductor haUiLcU in Chimp, as in man. Duvemoy £Dund it so in 
the Gorilla \ In An. it was different from that in man, and different 
on the two feet. On the left foot the proximal head rose with part 
of the Fl. br. d. and Abd. 5ti from the under Bur£eu^ of the Calca- 
neum, and joined the distal part of the second head (as was the case 
in the second head of the Fl. br. d). The second head rose by two 
fascicles, one from the internal, one from the external side of the. 
Scaphoid bone; these joined the tendon of the first head, and the 
common tendon was inserted into the extenxal side of the base of 
the first phalanx of the great toe. In the right foot the second head 
rose not from the Scaphoid bone, but from the deep &scia opposite 
it, and a third head was added, also from the deep &scia opposite 
the tarso-metatarsal articulation. 

Flexor brevis hcdlucia in Chimp, was very different fi'om that in 
man. Its inner head rose from several origius, principally from the 
Ecto-cuneiform bone, just where the second tendon of the Tib. p. 
was inserted. From this origin the internal portion (a) ran and 
fused with the lowest f in. of the tendon of the Abductor. This 
fusion is fonnd in man and in the Gorilla and Orang. The next 
portion {fi) was inserted into the internal sesamoid bone. An under- 
lying portion (y) rose from the Ento-cuneiform, and was inserted with 
the last portion (j3). BoUeston found the inner head prolonged by 
a tendinous expansion to the distal phalanx. The outer head, being 
that portion eventually inserted on the outer side of the tendon of 
the Flexor longns hallacis, wus thus arranged : the most internal 
portion (a) rose together with portions a and P of the inner head 
from the Ecto-cuneiform bone, and, crossing under the tendon of the 

^ In the Cebus and Innns nemestrinas it had two distinct heads, one from 
the Oaleanenm, the other from the plantar fascia. In the Orang it was inserted 
into the metatarsal as well as into the first phalanx. Yrolik describes two 
origins in Ghiup., one from the Ento-ooneiform, but also says it is as in 
man ; this second origin is probably part of the Flexor brevis, the fibres of 
which as in man fuse with the tendon of the Abductor haUnois. 



204 MR CHAMPNETS. 

Fl. 1. h., was inserted into the external sesamoid bone^ as is tbe 
case in man. The next portion (fi), having a similar origin and 
course, but larger, was inserted into the outer edge of the exteraal 
sesamoid bone, fusing at its insertion with some of the adjacent fibres 
of the Abd. h., which fusion is also seen in man. The deepest por- 
tion (y), consisting of 3 fascicles, rose from the Ento-cuneifurm with 
portion y of the inner head, and from the external side of the meta- 
tarsal of the Hallux in its whole length, and was inserted principally 
into the middle of the external sesamoid bone between the insertions 
of portions a and fi ; part however fused with portion p, and was 
inserted with it, and part ^ed directly with some adjacent fibres 
of the Adductor, and was inserted into the external edge of the 
external sesamoid bone^. In Ak. it rose by two distinct heads: the 
inner from the internal Cuneiform bone, with one of the palmar 
Interossei, was inserted into the internal sesamoid bone, and into the 
base of the first phalanx of the Hallux; the outer rose along the 
line of the middle metatarsal bone, from the deep plantar ligament 
and sheath of the tendon of the Flexor longus hallucis going to 
the Hallux, and fi-om a tendinous band of origin of the Interossei 
in that region. It was inserted into the external sesamoid bone, 
and the base of the first phalanx of the Hallux in common with 
the insertion of the Add. h. The identity of this muscle was proved 
by (1) its insertion, (2) its action, (3) its relation to the tendon of 
the FL 1. h., which separated its two halves. 

Addtictor haUucis in Chimp, was much larger than in man. Its 
principal origin was from the middle metatarsal in its whole length, 
and from an intermuscular septum between that and the fourth meta- 
tarsal. Its anterior and most transverse fascicle also rose from the 
distal end of the second as well as middle metatarsal, and its poste- 
rior or interior fascicle rose largely from the sheath of the Per. 1., 
and the superficial and internal aspects of the base of the second me- 
tatarsal. It was easily sepai*able into many fascicles, and might have 
been described as several muscles. Its general appearance was much 
more that of an Add. p. than of an Add. h., and it was stronger 
than the Add. p. It was inserted into the external side of the distal 
end of the metatarsal of the Hallux, some of its fibres fusing as in 



^ This last portion is called by Henle " Interoaseus volaris primusJ'* All the 
deep portions in the Chimp, also took origin from the sheath of the Per. 1. 
All the superficial factors of both heads also had origin from the sheath of the 
tendon of the Flexor longas hallucis. Yrolik describes the muscle as rising from 
the Ento-cnneiform, and forming a thin moscnlar fascicle. In the Gorilla and 
Orang Davemoy descrihes the whole muscle as rising from the Scaphoid and 
Ento-cuneiform. In the Orang Church found it rising from the Ento-cuneiform 
and plantar fasda, the external portion being inserted into the first phalanx, 
the internal into the metatarsal. In the Inuus nemestrinus the interior belly 
rose from the Ecto-cuneiform, and was inserted into the external sesamoid bone. 

The whole muscle was suppEed by the internal plantar division of the 
posterior tibial nerve. I could find no twig given from the deep branch of the 
external plantar to the external head, as sometimes in man (and as the analogy 
of the deep branch of the ulnar zftrve in the hand would lead one to expect) ; 
that branch seeming to lie at a deeper level also than this muscle. 



MTJSCLES AND NERVES OF A CHIMPANZEE AND ANUBIS. 205 

man with tlie adjacent part of the exterpal head of the Fl. l)r.* In 
An. it rose from the fascia and intermuscular septum between the 
metatarsals of the second and middle toes along rather more than 
their distal half; from a fascia, giving origin also to the plantar 
Interossei (thus showing its nature as an Interosseus), and to the 
external head of the Flexor brevis hallucis; also from the whole of 
the plantar surface of the base of the first phalanx of the second toe. 
It was inset-ted with the adjacent outer head of the Fl. br. h. into 
the outer side of the base of the firat phalanx of the great toe. Its 
origin thus differed widely from that in man. 

Abdtictor mimrrd digiti in Chimp, was large, rising from the 
whole of the anterior and plantar edge of the tuber Calcis, and from 
the plantar £iscia. Its external portion was inserted into the base 
of the 5th metatarsal. The next portion was inserted by a very 
delicate tendon into the external side of the bajse of the proximal 
phalanx of the little finger. The internal and main portion, which 
exhibited a penniform arrangement (a tendon which diminished from 
the origin of the muscle downwards, and disappeared before the in- 
sertion, lying on the middle of its plantar surface), developed one 
larger and two smaller tendons opposite the metatarso-phalangeal 
articulation, while other fibres were continued to the very insertion. 
These middle and internal portions were inserted just intei-nally to 
the first*. In An. it was as in man, and by its extensive connection 
with plantar fascia would give it tension and also flex the three 
exterior toes, owing to the frision of the plantar fascia with the 
tendons of the Fl. br. d. of those toes. In an adult An. there was 
also another muscle rising externally to it, and inserted into the 
base of the metatarsal of the little toe. 

Flexor brevis minimi digiti in Chimp., absent in RoUeston's 
Chimp., rose as in man. Its origin was complicated, by being shared 
by the palmar Interosseus of the little toe, and also by a muscle 
which certainly fulfilled the requirements of an Opponens. In An. 

^ Duvemoy, in the Gorilla, describes it as two mnscleB, " addacteur oblique'' 
and '* addnctenr transverse," according to the direction of the fibrea But no 
interral, such as that drawn by Dnvemoy (PL x.) in the Gorilla, could be seen 
in my Chihp., though its most transyerse fascicle could be divided from the 
rest, as could several other fascicles. Vrolik does the same, and says that the 
oblique portion comes from the Cuboid, the transverse from the fifth metatarsal. 
I could find neither origin. He says also that it consists in man of the same 
portions. Church, in the Orang, found a fascicle rising from a ligament 
" stretched from the head of the third digit to be inserted into the distal end of 
the metatarsal and proximal end of the first phalanx of the second," and 
inserted into nearly the whole length of the metatarsal of the Hallux. Cuvier 
also calls this portion '* adducteur transverse.'* 

' It is curious that the insertion into the base of the 5th metatarsal (that of 
the external portion) is correlated with the absence of the so-called Peroneus 
tertius in Chimp, and Cebns (Ghurdi, p. 17) ; but in the Innus nemestrinue 
this insertion coexists with the Peroneus tertius. In Chimp, it is weak, 
however, perhaps owing to the additional Extensor tendon given by the Per. 
br., and fusing with the Extensor tendon of the little toe. Vrolik found this 
muscle inserted into the second phalanx by a very fine tendon. Henle (p. 300, 
and Fig. 150, p. 296 a b q') describes an insertion into the tuberosity of the 
metatarsal as normal in man. 



206 MB CHAMPNEYS. 

it rose from the base of the 5th mletatarBal bone, and was inserted 
into the outer sesamoid bone and fibular side of the base of the firsii 
phalanx of the little toe. It was obviously an Interosseus. 

Opponens minimi digiti in Chimp, rose in common with the pre- 
ceding, and with the [mlmar Interosseus of the little toe, and was 
inserted into the external and plantar sur&uses of the 5th metatarsal 
in its whole length\ 

TrcmsveraaMs pedis was wanting in Chimp, as a separate muscle, 
as in the Orang, Cebus and Inuus nemestrinus, and occasionally in 
man ; but was represented by the transverse portion of the Adductor 
hallucis. 

Interossei in Chimp, were of the same number as in man, but 
abducted and adducted relatively to the middle digit as in the hand of 
man and Chimp., not relatively to the 2nd (index) digit as in the foot 
of man'. The first dorsal was much the largest, and had a broad origin 
from the base of met. i. as well from the side of met. i. like the Abd* 
indicis in hand. 

In An. the Dorsal Interossei were seven, and were inserted in the 
same manner as the plantar Interossei, and as in man. The internal 
(first and second) arose from the base of the metatarsal of the second 
toe, and were inserted one on each side of the second (index) toe. 
The third rose from the base of the metatarsal of the second (index) 
and middle toes, and was inserted into the internal (tibial) side of the 
middle toe. The fourth rose from the bases of the third and 
fourth metatarsals and was inserted into the external (fibular) side of 
the middle toe. The fifth rose with the fourth, but principally from 
the fourth metatarsal, and was inserted into the internal (tibial) side 
of the fourth toe. The sixth rose from the bases of the fourth 
and fifth metatarsals, and was inserted into the external (fibular) side 
of the fourth toe. The seventh rose from the base of the fifth 
metatarsal with the Flexor brevis minimi digiti, and was inserted 
into the internal (tibial) side of the fifth (little) toe. The dorsal 
Interossei alternately adducted and abducted from the middle toe. 

^ I can find no description of this mnsde in any of the anthropoid apes, 
but Henle and Huxley describe it as normal in man. It was found in a Chimp. 
by Bolleston. This muscle is described and figured by Dr Halford in the 
Macaque. He also describes a similar muscle in the Hallux, and says that 
" Professor Huxley has not shown that Table lY. does not apply to the foot of 
the GroriUa, Chucp., etc.*' Now, so far as Chimp, is concerned, I can answer 
that no muscle was inserted into the length of the metatarsal of its Hallux, 
though I can confirm Dr Halford's conjecture as to the presence of a similar 
muscle in the fifth metatarsal, as described by Huxley and Henle in man. Part 
of the flexor bre-vis hallucis rose from the whole length of the outer side of the 
first metatarsal, as I have described. 

' Duvemoy, who makes this comparison h propos of the Gorilla, refers it 
to the prepotence of the middle digit in the hand and foot of apes, and the 
second digit of the foot of man, as indicated by the superior length. This was 
also the case in the Macaque. The differences entailed by the point of abduction 
and adduction being the middle toe in the Anthropoid apes, the second in 
man, are as follows : — the middle toe in the Anthropoid apes, the Index in man 
has two dorsal (abductors), no plantar Interossei (adductors). The adaptation of 
the same general plan, the dorsal being abductors, the plantar adductors, is 
highly interesting. 



MUSCLES AND NEBVES OP A CHIMPANZEE AND ANUBia 207 

Table oi Dorsal Interossei in As, 

AddudUyra wliich were also partial flexora 

2tid adducted 2nd (index) toa 

5th addncted 4th toe. 

7th adducted 5th (little) toe. 
Ahductors. 

Ist abducted 2nd (index) toe. 

3rd abducted 3rd (middle) toe towards Tibia. 

4th abducted 3rd (middle) toe towards Fibula. 

6th abducted 4th toe. 

The 4th and 6th were also partial flexors. 

In An. the Plantar Interossei were three adducting to middle 
toe. Thej all rose from a common origin, viz. from the base of the 
middle met, and from a ligament stretched across the plantar space '. 

The external Plantar Interosseus ran to the base of the flrat 
phalanx and extensor tendon of the 5th toe, both on the tibial side. 
The second and first arose bj a common muscular slip, and afterwards 
divided and were inserted each bj a similar double insertion, the 
third on the tibial side of the fourth to6, the second on the fibular 
side of the second toe. They thus all adducted their toes towards 
the middle toe. 

In both feet a small slip of muscle ran from the interior (tibial) 
side of the first (internal) Plantar Interosseus (rising from the 
intermuscular septum between the dorsal and Plantar Interossei), and 
was inserted into the distal part of met. iii. This extra muscle points 
to the prepotence of the middle toe» as indicated also by its length. 
We find the same prepotence in Chihp., where the adduction and 
abduction are to and from this toe instead of the second, as in man'. 

^ This is mentioned by Church in the Innns nemeBtrinos and Cebns, and he 
does not seem to consider -these muscles as Interossei. It is, however, at least 
significant that they all addncted towards the middle toe which, as we have 
seen, is the prepotent digit in the foot of apes. He found the same arrange- 
ment in the hand. 

^ Though these two layers were distinct, and one lay more dorsally than the 
other, still, as Church remarks of the Inuus nemestrinus, there were no dorsal 
Interossei, as none had a dorsal origin, and also, but for the set which I have 
called plantar (as being more plantar than the other, and lying superficially to 
it), and which Church does not seem to consider Interossei, his account of them 
in the Inuus nemestrinus would tally with mine in An. Moreover, the real 
nature of the Fl. br. 5^ appears, for it is evidently an Interosseus. 

Duvemoy remarks that in the Gorilla, as I also found in Ghdcp., the 
dorsal Interossei are not so dorsally placed as in man. Moreover, this fact 
was plainly set forth in the hand of my Chivp., in which parts of the dorsal In- 
terossei were so far divaricated palmarly, as to be positively palmarly, and not 
at all dorsally, placed. In the lower monkeys, as Gebus, Inuus nemestrinus and 
in An., there are really no true dorsal Interossei, but two layers of Plantar, 
the more dorsally lying of which we may take, if we please, to represent the 
dorsal Interossei. The more plantarly placed resemble the Interossei of the 
Gamivora, as Church remarks. We therefore have an ascending series, from 
that ease where the dorsiJ Interossei are plantarly placed (represented by the 
Cebus and An., the more plantarly placed resembling those of the Camivora), to 



208 MB CHAMl»NEYS. 

AbdamincU Musdes, 

Rectus abdominis Jn Chimp, powerful, had two origios, as in 
man. The posterior wall of the sheath waa founded hy the aponeu- 
rosis of TransversKlis only, the anteiior bj those of the external and 
internal oblique, thus differing from the arrangement in man. It waa 
marked by four "inscriptiones tendinese " (as in Vrolik's) which went 
quite thi'ough the muscle to the sheath. 

In An. it became continuous with a fascia interposed between 
it and the inner layer of the Pectoral opposite the fifth rib, and was 
attached, as in man, to the cartilages of the fifth, sixth and seventh 
ribs. 

Pyramidcdia in both was absent as in Vrolik's specimen. 

External Oblique in Chiup. rose by seven digitations from 
ribs 5 to 11 inclusive, the two lowest interdigitating with Lat. 
d., the rest with Ser. m. It was inserted into the ant. sup. 
iUac spine for half an inch only, into Poupart's ligament and the 
linea semilunaris. Its fibres ended below at the level of one inch 
below the anterior superior iliac spine, and mesially opposite the line 
of the Rectus. In Ay. its first upper digitations were received not 
between those of the Ser. m., which failed to reach it, but of that 
special development of the Intercostals which has been already 
noticed. 

Internal Oblique in Chimp, resembled that in man. In An. 
it did not reach the last rib. 

Transversalis in both nearly as in man. The Fascia transversalis 
in Chimp, was better developed than in man, in coirelation with the 
increased strain on the abdominal muscles. 

Serves, 

The factors given according to Quain, and Flower's Plates. 

The innervation of the muscles closely resembled that in man, 
and did not call for remark except in the following particulai*s. 

Anterior Thoracic in Chimp., which supplied both Pectorals, seemed 
to receive supply from all the factors of the Brachial flexus, not 
only from C. V., C. VL, and C. VII. 

Nerve to the Subdaviua in Chimp, had no communication with 
the Phrenic. 

In Chimp, the Scalenus anticus was supplied by C. VIII., instead 
of CIV. 

Phrenic nerve was formed in Chimp, by C. IV. and V. ; in An. 
by C. IV. No connection could be found with the sympathetic in 
either. 

Middle cervical Ganglion was fused with the 3rd in both. 

Third cervical Ganglion in both was placed as in man at the 
root of the neck in an angle enclosed by the subclavian and vertebral 
arteries, which it more or less surrounded with plexuses. It sent 

the anthropoid apes, where these are more dorsal, aud thus to man, where they 
are more dorsal still. This dorsa migration of Interossei is very interesting. 



MUSCLES AND NERVES OF A CHIMPANZEE AND ANUBIS, 209 

twigs in company with the vertebral artery into the vertebral canal 
in the 7th cervical vertebra, another to join the 8th cervical nerve, 
another to join the recurrent Laryngeal nerve. 

Circun^fUx nerve in both was formed of C. V, VI, VII, instead 
of by C. V, VI, VII, and VIII. 

Posterior Tharotdo nerve (external respiratory of Bell), was formed 
in Chimp, by C. V. and VI. as in man. In An. by 0. VI., and 
VII. ; but the 5th cervical nerve, which supplied the Bh(Mnbpideus 
major, gave twigs to the serratns magnus, which thus received its 
supply from C. V. and VI (also as stated above from C. VII) though 
in a different manner from that in man. 

IntercosUhhumeral in An., unlike its representative in man and 
Chimp., pierced the lower part of the Scalenus post., which was in- 
serted in the 3rd and 4th ribs and not into the 2nd. 

Nerve to the Levator anguti Scapulce in Chimp, was C. IV, not 
C. III. 

Nerve to Khomboideus minor was C. III. in An., C. V. in Chimp. 
as in man. 

Supra-eeapular in Chimp., formed by C. V. alone instead of C. V. 
and C. VL ; nerve to Teres major in Chimp., was from the Sub- 
scapular, but from the circumflex in An. ; and this latter arrunge- 
inent has been noticed as an abnormality in the human subject by 
Prof. Turner*. 

Median in both passed under instead of superficial to the Bra- 
chial arteiy in the upper arm'. It communicated by its main branch 
with the ulnar nerve in Chimp.* but not in An. at about one-third of 
the distance down the forearm, and supplied the fingers simultaneously, 
and not from 2 main divisions. There waano supra-condyloid foramen. 

Ulnar nerve in Chimp, was small till it had received its &ctor from 
the median, when it doubled its size. It received a factor from the 
branch of the musculo-spiral nerve that supplied the Dorso-Epitroch- 
lien, afler which it gave some twigs to the inner head of the Tiicepa 
us it passed it, which possibly are derived from the branch of the 
mascalo-spiral joint membrane. 

MtuctUo-ciUanette in Chimp, passed through a cellular interval 
in the coraco-brachialit*, much more pronounced than in man« In An. 
it did not pierce the Coraco-brachialis but passed quite beneath it 
(dividing it from the Biceps), and nob superficial to it, as Wood* says 
is always the case in the Ist human variety, with which this arrange- 
ment otherwise corresponded. 

Gangliform enlargements over the back of the carpus, at the 
end of the posterior Interosseous nerve, and on the branch of the 

> Nat. Hist. Rev. Oct. 1864, p. 615. 

' Professor Turner Las noticed a similar arrangement several times in the 
human sdbject; and Prof. Humphry informs me that its ooomrrenoe is almost 
always associated with some abnormal disposition of one or other of the main 
arterial tmnks of the Umb, which is a point of some practical as well as morpho* 
logical importance. 

' I have jnst met with an example of this in a hxmian subject. 

* Camb. Joum. o/Anat. and Phyt. 1867, p. 45. 

VOL. VI. 14 



D. XIII. 



210 MR CHAMPNEYS. 

Ciicumflez going to the Teres minor were present in CHnfP. as in 
man. In An. they could not be distinguished. 

Flexor profundus digitorum in Chimp, was supplied by the 
anterior interosseous as well as by the main trunk of median and the 
ulnar. 

Flexor longus poUicie in Ohimp. was supplied from the main trunk 
of the Median as well as from its anterior interosseous branch. 

The general arrangement of the nerves of the lower limb and 
Lumbar and sacral plexuses was in Ohimp. Tery similar to that in 
man, but very different in composition, which was perhaps due to the 
fact that there were 13 instead of 12 dorsal vertebne. The differences 
in composition will be more readily seen by the following TaUe, 

Man Chimp. 

Ilio-Hypogastric, and ) t t 
Hio-Iniruinal. i 

Ctemtofcruri L. I. ll D. XIII. 

External cutaneous, L. II. III. D. XIII. L. I. 

Obturator, L. III. IV. D. XIII. L. I. II. 

Anterior crural, L. II. III. IV. D. XIII. L. I. II. III. 

Superior gluteal, L. IV. V. ; S. I. L. III. IV.; S. I. 

Sacral plexus, I* IV. V.; S. I. II. L. I. II. III. IV.; S. I. 

III. IV. II. 

Small sciatic, -L. IV. V.; S. I. IL L. lU. IV.; S. I. 

III. IV. 
Great sciatic, see Sacral plexus^ 

FeoM parvus in Chimp, was supplied by D. XIII. instead of 
L. II. In An. by L. H. Feoixe magnue by D. XIII., and anterior 
crural in Chimp, instead of by L. II. In An. the superior part (see 
description) by the ant. crural, the inferior by L. III. 

Fyriformia^ in An. but not Chimp, by the Superior gluteal, not as 
in man from the 2nd Sacral n. ObturcUor irU. in Chimp, supplied as 
in man from the sacral plexus, but the arrangement was different; 
for while in man one twig rising separately fi*om the plexus supplies 
it; a second, subsequently and separately arising, supplying the 
Gemellus superior; and a third, subsequently and separately rising, 
supplying the Gemellus inferior and Quadratus femoris; in Chimp. 
one nerve was given off from the sacral plexus, which bifurcated 
and sent one division to the Ob. int; the other division then gave off 
a twig to the superior edge of the Gemellus sup., then dived below 
the conjoined tendon of the Ob. int. and Gkmelli, supplying the 
Gemellus inf. and ended in the Quadratus femoris. (The two Obtu- 
rator muscles are never supplied by the same nerve.) In An. the 
same nerve supplied both Ob. int. and Cremel. inf.* 

^ It alBo differed from that in man in lying externally instead of iatemally 
to the Psoas magnns. 

' It will be observed in the arrimgement of the nerves of tite lower limb, as 
compared with that in man. that in many cases in which a^i^oent parts are 
BQpplied in man by nerves wnioh are given oft separately from a primaiy nerve> 
trunk, the same parts in these animalB are supplied by nerves |Jven off from a 



MUSCLES AND NEBYES OF A CHIMPANZEE AND ANUBIS. 211 

Coocygeua in Chihp. by the 3rd, and not the 4th and 5 th sacral 
nerves. The grecLt Sciatic nerve in Chimp, did not divide till the 
Popliteal space, but in An. almost as soon as it issued from the 
Pelvis. The point of bifurcation varies widely in man^ In both 
its distribution varied somewhat from that in man. In An. the 
proximal end of the long head of the Biceps (the short head being 
absent) was supplied by the int. pop. n. the distal part by the ext. 
pop. The Semitendinosus and Semimembranosus were supplied to- 
gether by a coitimon branch of the great sciatie, instesul of by separate 
branches, anothev instance of lowness of Integration. In Chimp. 
a separate trunk from the great Sciatic high up after a course «f 
4 in. gave off (1) a branch to the origin of the Semitendinosus; 
(2) a branch which supplied the upper third of the long head 
of the Biceps; (3) the remainder split into branches which sap* 
plied the prpximiai part of the Add. m., the mesial part of the 
Semitend. and Semimemb., and the separate external division of the 
Add. m. (noticed under Uiat muscle). The short head of the Bi* 
ceps was supplied by two twigs separately rising from the main 
trunk of the great Sciatic nerve, nearly opposite the middle of the 
thigh. Another instance of want of Integration; but it is curious 
that the anthropoid Chimp, fiunishes a better instance of it than the 
low An. 

JSartorius supplied in Chimp, by the main branch, and also (un* 
like that in man) in its lower part by the Internal Saphenous.- 
Fectineus in Chimp, supplied by a somewhat large branch from the 
Ant. cr. n., and by a very small branch from the ob. In An. by Ant. 
cr. n. only. (In man the twig from the obturator is inconstant.) 

seoandary braneh together; that is to say, in man the primary nerre-tnmk has 
absorbed the seoondary trunks into itself. This is a ^ood instance of the 
advance in "Integration" in man, as compared with animals less highly or* 
ganised. 

1 Qaain, Vol. n. p. 676. Turner (Nat. HUt. Rev. Oct. 1864, p. 616) says 
that snoh eariy duplicity, whan it oconrs in man, almost always coexists with 
dnpUoiiy of the Pynformis. 



14—2 



PARTIAL DEFICIENCY OF THE TENDON OF THE 
LONG FLEXOR OF THE THUMB. By W. W. Wag- 
8TAFFE, B.A.^ F.R.G.S., LecttMrer on Anatomy <U St Thamaa'a 
HoipUal, 

PECULiAaiTiES in the arrangement of the extensor tendons to the 
fingers are of common occarrence, but it is rare to find any great 
deviation from the nsual plan of arrangement in the flexor tendons. 
I am, therefore^ led to record an instance of abnormality in the long 
flexor tendon of the thumb, which came under my notice in the 
dissecting room at St Thomas's Hospital early in the beginning of 
last year. 

During the usual demonstrations my attention was called to what 
was thought to be a case of deficiency of the long flexor tendon of 
the thumb (left), and upon careful examination the following con- 
dition of parts was found. A muscle (t) occupying the usual posi- 
tion of the flexor longus pollids arose from the nuiius and inter- 
osseous membrane, and its tendon passed downwards in its usual 
position under the anterior annular ligament and the superficial 
flexor tendons. Instead, however, of passing between the two heads 
of the flexor brevis to the thumb, the chief jjart of the tendon (1) 
joined the index tendon of the flexor profundus digitorum (Fl. p.) : a 
second portion (2) spread out upon the carpus and attached itself to 
the ligaments covering the carpal bones in front : and a third 
part (3) left the tendon rather higher up, and was inserted by a 
well defined triangular expansion into tlie outer snr&ce of the 
outer or superficial head of the flexor brevis poUicis. 

On the thumb itself a well-marked tendon (st) lay on the palmar 
surfibce of the first and second phalanges in the position usually 
occupied by the digital portion of the flexor longus pollicis tendon. 
This peculiar tendon was attached by two distinct slips to the head 
of the so-called metacarpal bone of the thumb. These two slips 
were connected one with each side of the head of the bone. From 
these proximal attachments it passed forwards, lying free upon the 
first phalanx, and was slightly spread out to be inserted as usual 
into the base of the ungual phalanx. A reflected portion represent- 
ing the ligamentum breve (l. b.) was connected as usual with the 
capsule of the neighbouring joint and the head of the proximal 
phalanx. 

The Nerves were also peculiar in the thumb. Two digital 
branches as usual came from the median, but about an inch from 
their origin they each presented enlargements — on the outer nerve 
two^ on the inner one — in size about equal to a millet seed. Beyond 
these enlargements the outer nerve could be traced onwards to its 
ordinary distribution, but the inner nerve appeared atrophied and 
was lost in fibrous tissue opposite the middle of the first phalanx. 

The arteries were smaller than osoaL 



PARTIAL DEnCIENCT W FLEXOR OF THUBIB. 213 

In this case, therefore, we have a muscle arising like the flexor 
longus poliicisy but inserted into the deep flexor tendon to the index 




finger, and connected also with the fibrous structures about the 
wrist ; we have a tendon resembling that of the flexor longus pol* 
licis, but only stretching between the head of the metacarpal bone 
and the ungual phalanx ; and associated with these muscular pecu- 
liarities about the thumb we have peculiarities in the appearance 
of the digital nerves and ressels. 

That these peculiarities were developmental appeared beyond 
doubt) for there was no evidence either in the existence of scar^ or in 
the matting together of neighbouring tissues, that mechanical injury 
or disease had been the cause. It would seem that an error of develop- 
ment had occurred; that from some cause the intermediate portion 
had not been developed; that consequent upon this the free ends had 
attached themselves to neighbouring parts — one to the head of 
the metacarpal bone, the other to the deep flexor tendon for the 
index finger and the cari)us. And the condition of the digital 
nerves and vessels would agree with, although it would not neces- 
sarily require, the supposition that these peculiarities were the result 
of fiiulfy development. 

It may be asked what movement was obtained in the thumb, 
and I can only judge by experiment made upon it after it came 
under my notice. No fiexion of the ungual phalanx was obtained by 
traction upon muscles. The ordinary movements of the proximal 
phalanx were obtained. Traction upon the muscle which occupied 
the place of the flexor longus pollicis produced flexion of the index 
flnger, with some flexion of the wrist. It did not seem that any 
means were provided for flexion of the tip of the thumb by connec- 
tion between the flexor brevis and the slips of tendon lying alongside 



S14 ME WAQSTUPE. 

its insertion. It moat be concladed, therefor^ that the ongaal pha- 
luix of the thamb in this otse was not capable of inovemetit at the 
will of its owner; and as no histoi; oonld be obtained of the old 
woman's habits, owing to her being an nnclaimed snbject, I am 
unable to gire the historical evidence which might be intOTesting. 



DESCRIPTION OF Alf ACCESSORT MUSCLE IN CON- 
NECTION WITH THE POPLITEUS. Bjr "W. W. "Wao- 
ffTAFPZ, RA., F.R.C.S., Zeclurer on Anatomtf at St Tkomat't 
Soapital. 

In dissecting the popliteal space of the left leg of a subject last 
session, an onnsual muscle was found in connection with the 
pop] i tens. 

The outer head of the gastrocnemius possessed a lather lu^ 
sesamoid (S) bone close to ita attachment to the femnr. From the 
inner side of thia sesamoid bone arose a rather strong tendinous 
structure, distinct &om the gastrocneinius proper, and sooa expanding 
into a well-marked muscle, which passed downwards to the tibia. 
The fibres were directed inwards as well as downwards, and lay 
superficially to those of the popliteus {P), ultimately becoming at- 
tached to the inner edge of the tibia as ^ as the oblique liae, and 
also blending with the fibres of like pofditeos. The po{^teal &G<ua 



A MUSCLE ACCESSORY TO THE POPLITEUS. 215 

WB8 spread over that portion of the muscle which was in contact 
with the popliteus. 

It is worthy of note that the plantaris was absent, and that the 
general muscular derelopment was not excessive ; and also that no 
similar abnormality was found in the opposite limb. 

I have searched without saooees far the record of any similar 
abnormality in the human subject^ and I do not find in comparative 
myology much that will throw light on such a variation. I know 
of no instance in which the post-condyloid or gastrocnemial sesamoid 
bone gives origin to portion of the popliteus; but in the Anteater 
Professor Humphry describes the outer head of the gastrocnemius 
as arising from the popliteal sesamoid. {Jaum. ofAncU, iv. 59.) 

The use of the muscle in the present instance was obvious. It 
acted as a flexor of the leg upon tiie thigh, and assisted slightly in 
rotating the tibia in the first movement of flexion. 



UTERINE CONTRACTIONS DURING PREGNANCY. 

Dr J. Braxton Hioks, F.R.S., read a paper at the Obstetrical 
Society of London, on 4 Oct, in which he pointed out a ieict which 
he considered had not been noticed before ; namely, that the uterus 
not only daring the last month of pregnancy, but from at least the 
third month, contracted and relaxed frequently. These contractions 
oocnired every 5 to 20 minutes generally, although sometimes the 
intervals were longer. They lasted about 3 or 4 minutes, but in the 
case of a diseased ovum the contractions were frequently longer, and 
sometimes almost continuous. They took place even in cases of 
vesicular mole (Hydatiniform degeneration of the Chorion). Dr 
Hicks had only in one case noticed the absence of these contractions, 
viz. in a case of Paraplegia, at least while it was under his observa- 
tion for two months. When the uterus was retroverted in the early 
months of pregnancy the uterine body was more readily under obser- 
vation and its state easily noticed These contractions are not owing 
to the external irritation during examination, but as frequently as 
not the uterus is found upon first examination to be hard, and then 
to relax. After describing the physical state of the organ during 
these conditions, Dr Hicks alluded to the value of these contractions 
physiologically. He thought at least two advantages were derived 
from them : one to supplement the heart impulse in a part so far 
removed from its efleots : the other to assist the ultimate disposition 
of the foBtus. After this he discussed at length the assistance to 
diagnosis these frequent contracti(Mi8 gave the practitioner. 



ANOMALIES OF ARRANGEMENT. By D. Emblbtoh, M.D. 

MUBCULAB. 

The anomalies here given were observed in the Dissecting Room 
of the then Newcastle-upon-Tyne School of Medicine and Surgery, 
during the dissection of the body of a muscular sailor in 1842. 

The Biceps Flexor CvJbUi had, as is not very uncommon, three 
heads, two of which arose in the ordinary way, the third took its 
Heshy origin from the inner side of the humerus along the ridge 
extending upwards from the inner condyle between the contiguous 
edges of the Brachial is Anticus, and the Triceps Extensor, and in 
front of and below the insertion of the Coraco-brachialis. It was 
about two inches in length of origin, was thick, and resembled, 
except that it was smaller, the femond head of the Biceps Flexor 
Cruris. It joined the common tendon of the muscle at the inner 
side and back part, at the bend of the elbow. 

On both sides there was no distinct Flexor hrevis minimi Digiti, 
The FcUm^aris brerns was very largely developed. The outer edge of 
the Supinator Radii longus just where the muscle ends in the tendon 
was inserted into the fascia of the arm covering the extensor muscles. 
The Fronator teres had no origin from the coronoid process of the 
ulna, arising only firom the inner condyle of the humerus. The Infra- 
spinattu and Teres minor were united as one muscle. The Teres 
major very thick and strong. 

In the left arm there was no Falmaris lortgus, though in the 
right it was present and normal. The Flexor Carpi radialis and the 
Flexor Carpi tilnaris were both attached to the Palmar Fascia, the 
former more directly and more extensively than the latter, which 
however was more connected with the fasda than is usual when the 
Palmaris longus is present, but the Flexor Carpi radialis chiefly 
supplied the place of the Falmaris longtis. 

There were no Fyramidales abdominis. The Fsoas pa/rous was 
absent from both sides. The Cremasters arose in the mode described 
by Gabriel, Fallopius, and even by Galen, long before M. Jules Cloquet 
had dissected them and claimed the discovery of their airangement. 

In the Ferinieum were two Vran^swrsi muscles on each side. 

In the deep posterior region of each leg was an additional small 
muscle of an interesting character. It arose in each leg, fleshy, from 
about the inferior third of the posterior surfietce of the flbula, but not 
quite so far down as the malleolus extemus ; the fleshy fibres passed 
backwards to a slender tendon, forming a simple and short, but thick 
and strong penniform muscle. The slender tendon passed down 
towards the sole of the foot, between the inner ancle and the heel, 
behind the Flexor FoUicis longtis, and therefore behind all the other 
parts lying in that .region. In the sole it was found between the 
tendon of the Flexor longtis digiiorum communis and that of the 
Flexor Umgus Follicis, where it divided into two slips, one being 



DR EHBLETOK. ANOMALIES OF ARRANGEMENT. 217 

firmly attached to the innermost division of the former tendon, and 
the other to the outer side of the latter tendon. This muscle was 
thei'efore an assistant flexor of all the toes, aiding both the common 
flexor and the special flexor of the great toe, and supplanting the 
nsnal arrangement by which these two muscles are combined in 
action by a strong oflset from the tendon of the latter passing to 
unite with the tendons of the former. (The muscle above described 
must be rather a rare anomaly, and I do not find it mentioned in any 
of the books to which I have access.) The Flexor ctccessoritta was 
well developed* The Feronetts tertitts ended in two small ten- 
dons which passed, the inner to near the base of the fourth meta- 
tarsal bone, the other to the ligament connecting the base of the 
fourth to that of the fifth metatarsal. 

In another subject, the Oemdltis superior, on the right side, was 
absent, the Obturator inteiiius being stronger than usual, and to its 
tendon that of the Gemellus inferior was scarcely at all adherent. 



Arterial. 

In Session 1845 — 46. In a boy of 17 years the arrangements 
at the arch of the aorta were as follows : there were only two pri- 
mary branches from the aroh, viz. the Brachio-cephalic and the left 
subclavian, the former immediately divided into two secondary 
branches, the Brachio-cephalio proper and the left common carotid, 
the latter, the left subclavian, pursued its usual course, whilst the 
left carotid crossed over directly to its proper position. 

The heart was altogether higher placed in the thoitkx than usual, 
its apex pointing to the interval between the fourth and fifth ribs, 
near to the costal end of their cartilages. On the left side were two 
Ascending Phar3mgeal Arteries ; the superior thyroid was given off 
from the common carotid two or three lines below its bifurcation ; 
the lingual and the fieunal came off by a common trunk, and there 
were two branches from the external carotid just above the bifur- 
cation, to the pharynx. The posterior amis came from the external 
carotid. There were two renal arteries on each side. 



Nervous. 

In a foetus dissected in 1840, the Chorda Tympani Nerve, after 
issuing from the fissure of Glaser, applied itself, not to the gustatory, 
but to the inferior dental nerve, and leaving it at a line or two above 
the orifice of the dental canal, passed on in the direction of the sub- 
maxillary gland, and Before joining the submaxillary ganglion, sent a 
branch of communication to the gustatory nerve, and twigs to the 
submaxillary and sublingual glands. 

Mai/ 27, 1871. 



REPORT ON THE PROGRESS OF PHYSIOLOGY*. By 
T. Laudeb Bbunton, M.D. D. Sc.^ Jovnl Lecturer on Materia 
Medloa and TherapeuticSf St. Bartholomeu^e ff capital, and David 
Febbieb, M.A., M.D., Demanatrator of FraeticcU Physiology ^ 
King'' 8 College, London, 

Nervous System, 

FuKcnoN OF THE Cebebbal PEDUKCI.B8. — ^In liifl eoqierimeats on 
this subject, Afismasieff (fTien. Med. Wockenseh, pp. 137, 153, 169, and 
185) divided one or both pednncles through a hole in the temporal 
bone. In eonsequenoe of the transitoiy irritation which the section 
produced, the animal immediately afterwards drew itself together, the 
head was inclined to that aide on which the peduncle had been 
flivided, the pupils became contracted espedaliy on that side, and the 
arteres of the ears also contracted, but their contraction was more 
marked in the ear of the opposite sida In six seconds after the 
operation all the above-mentioned effects were succeeded by their 
opposites. The irritation also produced increased flow of tears and 
saliva, and twitchings of the extremities on the side opposite the 
section, all of which lasted for half an hour. Section of one peduncle 
between the pons and tuber cinereum poroduoed paralysis of the muscles 
of the extremities on the opposite side, and <^ those of the back and 
neck on the same sida The amount ci paraljrsis increased wiih time. 

The character of the man^ and pointing movements made by 
the animal after the operation dianged through time, and in two or 
three weeks it was again able to run straight fbrwiuxL A section 
opposite the tuber cinereum, or in front of it, produced paralysis of 
the muscles of the back and neck as well as of the extremities on the 
opposite side, shewing that the nerves from the dorsal and cervical 
muscles cross opposite the tuber, whil^ those of the extremities cross 
lower down in the peduncle. Section of the peduncle causes c<Nn* 
plete paralysis of the oculo-motoiius on the same side and imperfect 
paralysis of the facial nerve on the opposite sida It also produces 
diminished sensibility of the body and head, and contraction of the 
arteries, which lasts for 10 or 15 days, and is more marked in .the ear 
on the same side as the section, than on the opposite one. 

At the moment of section the blood-pressure is increased and the 
pulse slowed, in 30 or 40 minutes after it the temperature of the 
body becomes lowered from 1 '5 to 2^ C. Division of both peduncles 
destroys the power of voluntarily relaxing or tightening the sphincter 
ani as well as of relaxing the constrictor urethne. It does not affect 
the movements of the bladder. 

Centbal Ibbadiatiok of the impulse of the Will, — ^Nothnagel 
(Arch,/. PsyckUU, und Nervenkrank. iii. 214 — 218) describes a case 
of what he terms central irradiation of the impulse of the will. A 

^ PhjBiologioal papers to be sent to Dr Bnmton or Dr Ferrier, 28, Somerset 
Street, Portman Square, London, W. 

[The Reports by Prof. Turner and Dr Fraser are unavoidably postponed till 
next number.] 



REFOKT ON THB PROGBESS OF PHTSIOLOOY. 219 

|>atient) after an attack of typhus, suffered from a carious affisction of 
the left leg and right arm. Only after great trouble and the lapse of 
several minutes could the patient flex the right arm when he wished. 
The biceps became hard and contracted, and at the same moment the 
triceps also contracted and offered strong resistance to the flexor 
muscles. 

These phenomena only occurred on slow or moderately quick 
movements. The flexors and extensors of the left leg were simi- 
lariy affected, but to a less extent. From the &ct that these ano- 
malies coidd not be induced by mechanical irritation, and that the 
contraction of both muscles began at the same time, and also from 
the circumstance that when one of the muscles was irritated directly, 
it contracted, while its antagonist remained at rest, the author con- 
cludes that this was an example of central perversion of innervatioo, 
not of reflex origin. 

iRBITABILrrT OF TBK ObMTBIFETAL FiBRES m THB SPINAL CoRD. 

— ^Ludwig and Dittmar (Lu(iung*8 Arbeiteny 1B70, p. 4) use the rise 
of blood-pressure which occurs in curarized animals when a sensory 
nerve is irritated, as a test of the conduction of sensory impressions 
to the brain, and consider it to be a much more delicate index than 
the movements or struggles of the animal. They find that the blood'* 
pressure rises in proportion to the amount of irritation, when other 
disturbing factors, such as exhaustion or changes in the respiration, 
or pulse rate, &c. are excluded, and each irritation has the same dura- 
tion. The dura mater is one of the most sensitive parts in the body, 
and the mere opening of its sac causes the blood-pressure to rise as 
much as the operation of separating the posterior from the anterior 
columns of the cord. When the cord was exposed and divided, and 
the nerve^oots cot or torn, a slight irritation of the central end lasting 
for. several seconds, .such as rubbing it with a blunt needle, or the 
application of a very weak galvanic current, caused a considerable 
rise in .pressure. A 5 per cent, solution of caustic potash also caused 
a risoi' Momentary irritations, mechanical, chemical or. electrical, 
however powerful, luui no effect; caustic potash having no action, and 
sparks from an induction coil causing no rise even when repeated 
every three seconds. When the cord was divide into anterior and 
posterior halves irritation of both caused a rise, but that produced by 
irritation of the anterior half was the greater, peirhaps fix>n^ the ante- 
rior roots or dura mater being also irritated. When the cord was 
raised from its bed and divided into parts irritation of the anterior 
columns, or of the grey substance, had no effect ; irritation of i^e 
lateral columns caused a slight rise, and irritation of the po6teri<Mr 
columns a rise of pressure. 

Irritation of the peripheral stump of the cord caused a rise some^ 
times greater and sometimes less than the reflex rise, and, at the 
same time, a slowing of the pulse. The effect of irritation of the 
central stump of the cord was inconstant l^ough quickening was 
often observed. The influ^nce of the point irritated, or of the nature 
OP strength of the irritation on the pulse, is not constant. For 



220 DR BEUKTON AM) DR FERRIER. 

an explanation of this we mast refer to the original. The rise of 
pressure oconrs after the cerebrum has been separated from the 
medulla. This shews that it maj take place without the animal feel- 
iog. When the cord was irritated after separation from the cere- 
brum no rise took place at firsts but this was due to an accumulation 
of blood in the spinal canal, and when the blood was removed a rise 
occurred as before. Although irritation of a sensory nerve may cause 
reflex independently of sensation, there is no reason for believing that 
there are special vaso-exdto-motor fibres distinct from sensory one^ 

Time requisxd to oommukicatb Impressions to the Sensobium, 
AND the Revebse. — As a means of measuring the time necessary for 
this, T. C. Mendenhall (American Journal of SdvnM cund ArU^ 3rd 
Ser. il), made use of an astronomical ohronographi which moved a slip 
of paper at the rate of 1^ inches per second. The seconds were marked 
on this by means of electridly. The person experimented on was seated 
before an opening at which a red or white card could be exhibited, 
and his flnger rested on a knob which at the slightest pressure dosed 
a current and produced a dark spot on the moving paper. This knob 
he was directed to press as soon as he saw the card appear at the 
opening. The exact instant at which it came before the opening was 
noted by electricity on the paper, and by measuring the distance 
between this point and that produced by pressure on the knob the 
time requisite for the impression to travel up to the sensorium and 
back through the motor nerves to the finger could be easily ascer- 
tained. By causing cards of different colour or shape to appear, and 
desiring the person to indicate their character, by pressing on one or 
other of two knobs, the time requisite for dedding between them was 
ascertained. 

The time required to respond to the appearance of a white card 

was ... ... ... ... 0*292 seconds. 

... an dectric spark 0*203 

... ... a sound 0*138 ••• 

touch on the forehead 0*107 

... touch on the hand 0*177 •.. 

when required to dedde between white and red 0*443 

a circle and a square 0*494 

... the tones C and E 0*336 

C and C above 0*428 

Bensobt Paths in the Spinal Cord. — Miescher, under Ludwig's 
direction (Ludwig^s Arbeiien^ 1870, p. 172), performed a series of 
experiments in order to determine what parts of the spinal cord have 
the power of conducting impresdons. Like Dittmar, he used the 
rise of blood-pressure, which occurs on irritation of a sensory nerve, 
as an index of the conduction of the impression to the brain. The 
experiments were made by irritating the sciatic nerve and comparing 
the rise in blood-pressure which ensued before and after the division 
of parts of the spinal cord. The cord was reached by cutting out a 
part of the laminae between two spinous processes. In order to limit 



REPORT ON THE PROGRESS OF PHYSIOLOGY. S2l 

the extent of the section more aocuratelji a Bmall knife 8 mm. long 
and 2 mm. broad, was driven with its flat side parallel to the longitu- 
dinal fibres of the cord quite through it^ till it became firmly 
fixed in the bone in front. The part of the cord lying on one or 
other side of it was thus completely protected, so that either part 
could be entirely destroyed without fear of injuring the other. When 
one or both lateral columns were left and the rest of the cord com- 
pletely divided, irritation of the sciatic nerve caused the blood-pres- 
sure to rise just as before. The irritation in this case was not con- 
ducted by means of small portions of grey matter still adhering to 
the column, since the result was quite as well marked when the 
column was absolutely free from them. When both lateral columns 
were cut there was no reaction, although the rest of the cord re- 
mained intact. 

This shews that the impressions on the sciatic nerve which pro- 
duce reflex rise of blood-pressure are conducted chiefly, if not entirely, 
by the white lateral columns of the cord. When one lateral column 
was preserved, rise of pressure followed irritation of either sciatic, 
but it was higher when the opposite nerve was irritated, shewiDg 
that in that part of the cord which lies between the last thoracic and 
third lumber vertebra, impressions from the left sciatic are chiefly 
conducted by the right lateral column, and to a less extent also in the 
left, and vice versa. The crossing of the conducting paths takes place 
gradually, and those which enter the cord low down run in its most 
external part, while those which enter higher up run nearer to the 
middle lina No hyper-SBstheeia was observed in the hinder extre- 
mities after partial section of the cord. 

Chobeiforu Movememts in Doos, akd Action of Electricitt 
ON Keflex Movements. — Legros and Onimus (Journal de VAnat, et 
Phys.y 1870, p. 403) find that the choreiform movements in dogs vary 
in character and are generally not coincident with the cardiac pulsa- 
tions. They have their cause in the grey substance of the spinal coi*d, 
but may be modified by the action of the brain. They are lessened 
by division of the posterior roots of spinal nerves. Their disappear- 
ance daring sleep shews that the spinal cord sleeps as well as the 
brain. They are lessened by chloral. The cord presents no patholo- 
gical appearances except slight congestion. A constant electrical 
current passing upwards through the cord causes irritation, increases 
reflex action, and does not remove the tetanus produced by strychnia 
and picrotoxin. A current passing downwards produces inhibition 
of reflex action and at once stops the tetanus produced by these 
poisons. The choreiform movements are increased by an ascend- 
ing and diminished by a descending current. When tiie current is 
opened an opposite eflect is produced. 

Influence of the Posterioe Roots of Spinal Neeves on the 
Iesitability of the Anterior Boora — Steinmann and Cyon (Bull, de 
VAcad, imp. dee JScieneea de St. Petershourgj vu. Dec. 1870) find that 
when the posterior roots of spinal nerves in frogs are quickly divided 
without pain to the animal, irritation of the anterior roots produces 



222 BB BEUNTON ASD DR FERRIER. 

a weaker contraction of tbe mnsdeB than befinra If pain be jMro^ 
duced by the division, the oontractiona immediatelj after it are 
stronger than before, and then they become weaker. This is due to 
the direct irritation of the anterior root being strengthened by that 
produced by the pain. After the posterior roots were divided the 
muscles became extended more quickly and to a greater extent after 
a contraction than when the nerves were intact. If a weight were 
attached to the- muscle, it became longer when the posterior roots 
were divided. They regard this as a proof that the musdes are natu- 
rally in a state of slight tonic contraction, due to excitations conveyed 
to their motor nerves through the posterior roots. 

■ 

Trophic Lesions. — Fischer {Berlin, Klin. Wochenachr. 1871, 
No. 13) describes the phenomena, viz. oedema, impaired nutrition of 
the nails, skin eruptions, and lastly, neuroparalytic ulcerations of the 
fingers and toes, which result on lesions of the nerves of the extrer- 
mities. The temperature of the paralysed parts is at first higher, 
and afterwards lower than normal ; generally there is much more 
copious perspiration. The author rejects the idea that these pheno- 
mena are merely traumatic and due to loss of sensibility. He finds 
that the sensibility may be quite normal. He agrees with Samuel in 
attributing the effects to paralysis of special trophic nerves. 

Schiefferdecker {Berlin. Klin, Wochenschr. 1871, No. 14) records 
several cases of trophic disturbances following lesions of nerves. The 
phenomena were atrophy of the muscles, thickening of the skin, 
desquamation, iiTegular growth of hair and nails, and increased per- 
spiration. Some of these phenomena he thinks are undoubtedly due 
to paralysis of special trophic nerves. 



Electric Stimulation op Nerve. — J.K6nig(F&n. Acad. Sitsi>er. 
LXii. 2 Abth. 537 — 46) has put Dubois Keymond's law of nerve sti- 
mulation to the test of fresh experiments. As a stimulant he em- 
ployed the irritation caused by closure of the descending current. 
From his experiments it appeai'ed that no contraction occurs when 
the duration of the current is very shorty and that from the minimum, 
increased duration of the current causes the height of contraction to 
rise at first rapidly and then more slowly, until, with a duration of 
the current from 0*025 — 0*018 sec, the curve blends with the asymp- 
tote which represents the height obtained by prolonged closure of the 
same current. Lamansky's view is thus confirmed, that the so-called 
super-maximum contractions arise from the summation of two stimuli 
He agrees with Dubois, that for weak currents of short duration 
the excitation depends chiefly on the amount of variation in the in- 
tensity of the current from one moment to the other. In accord- 
ance with the observations of Brucke, it would appear that the elec- 
tric current, in order to produce the molecular change corresponding 
to excitation of a nerve, must have a duration of at least 0*0015 sec. 
Even then the irritation does not occur instantaneously, but occupies 
a certain time with gradually diminishing intensity. The au^or 
gives a formula which includes these various &ctors. If the nerve is 
cooled down to 0', a duration of the current of 0*02 see. is necessary 



REPORT ON THE PR00RB3S OF PHYSIQL06T. 22S 

tn order to prodnoe irritation of the nerve, aod at this temperature 
the differences in the amount of irritation seen at ordinary temper^ 
aturee are no longer obeerved. 

Negative Oscillation op the Nerve-current. — Schiff {Pflil- 
gef^sArchiv, iv.) adduces further evidence in support of his view, that 
the negative oscillation which is observed when a nerve is irritated is 
dependent on an " irritation-current," and not on the natural current 
of the nerve when in the state of rest. 

Elbotro-motor Properties of EifRRTONic Nerve and Muscle. — 
'^- Yalentin (ZeUschri/lf. Biol. vii. p. 105) finds that before the 8th or 

10th day the nerves and muscles in the chick are not sufficiently 
differentiated to allow of a satisfeustory examination. On the 11th 
day the muscles have the power of contraction, but both they and 
the nerves are far from being completely developed. Nevertheless, 
they both sliew the same currents^Ahat they do in their perfect state, 
as well as the negative oscillation during action. 

Development of Heat in Nerves and Nerve-centres. — Schiff 
{Arch, de Fhys. Norm, et Path. ii. lii.) has investigated the question 
of development of heat in nerves and nerve-centres consequent on 
sensorial and sensory impressions. His experiments were made by 
means of a very delicate thermo-electric pile of antimony and bismuth. 
The development of heat in nerves was less marked the &rther from 
the point of irritation. Schiff explains this by diminution of the irri- 
tation during transmission. In the case of the bi'ain he found that 
during life there was a distinct development of heat in the hemi- 
spheres on the occurrence of a peripheric stimulus, such as pressure. 
The deviation of the needle indicated unequal development of heat in 
< the hemispheres. The development of heat is greatest in the middle 

of the hemispheres. 

Schiff {Ffluger^B Arehiv, rv.) irritated from the cord the sciatic 
nerves of both sides, one of which had been cut at its junction with 
the gastrocnemius muscle, and expected to find a development of heat 
in the cut nerve. He found, however, tiiat there was no difference 
between the two, or even a development of heat rather in the uncut 
nerve. He offers no explanation of the phenomenon. 

Bernstein, IiweetigaHona on the SHrmdation of Nerve ondMumdef 
Heidelberg, 1871. 

Wundt^ Mechamam of the Nerves <md Nerve-eenJtreSj Erlangen, 
1871. 

Eulenburg, Dieeaeee of the Nervous System considered Physio^ 
logically, Berlin, 1871. 

Vision, 

Time rbquisxtb for Yisual Perception. — haxt(PJl4tger'sArGhii^, 
tv. 325 — 36), under the direction of Helmholtz, has made some very 
interesting observations on the time required for a visual impression 
to affect consciousnesB, and on the extent of the percept in visual 
impressions of given duration. 



224i DR BRUNTON AND DR FERRIER. 

The experiments of Helmholts and Exner had shewn that when 
a number of black letters are printed on a white ground, one, or two, 
or more of these can be distinguished according to the duration of the 
retinal impression. These results form the starting point of Baxt's 
experiments. By an ingenious apparatus, the image of a number of 
black figures printed on a white ground could be allowed to fall on 
the retina for any fraction of a second, at the end of which time the 
impression was succeeded by a very much stronger stimulus. In this 
way could be estimated the time which simple impressions require in 
order to affect consciousness. For the time requisite for peroejition 
is that time after which a weaker impression can no longer be effaced 
by a stronger. 

Within certain limits, it was found that the time requisite for the 
first impression was independent of the degree of Dlumination of the 
object. The experiments shewed that consciousness is able only 
gradually to elaborate a griven impression, and that between the 
direct stimulus and the perception in consciousness one-twentieth of 
a second intervenes. 

It appeared also, although the experiments could not be carried 
&r, on account of the £sktigue to the eye, that a very strong second 
stimulus is able to efface the first eyen after a longer interval. In re- 
gard to the extent of the percept, the results shewed that in all degrees 
of intensity of the impression a much longer time is necessary to per- 
ceive small objects (axich as intervals of ^pace) than large ones. Thus 
with a duration of tne impi'ession for 0*0005 second, and good illumi- 
nation, large letters could be distinctly perceived, while smaller ones 
of the same form were very indistinct or quite imperceptible. Further 
experiments in reference to Fechner*s psychophysical law, ^* that dif- 
ferences of illumination are more easily perceived the less the general 
degree of illumination," proved, that in proportion to the difference 
in the illumination of two different objects the time, requisite for per- 
ception diminishes. 

Time Necessary for Vision. — Ogden N. Bood (American 
JoumcU of Seienee and Arty 3rd Ser. ii. 159) finds that the 
letters on a printed page may be plainly seen, and if a polariscope 
be used, the cross and rings round the axes of crystals can be ob- 
served with all their peculiarities, and errors in the azimuth of the 
analyzing prism noticed by an electric spark, whose duration is only 
forty billionths of a second. 

Subjective optical phenomena can also be produced, such as 
Loewe's rings, using cobalt glass, and the radiating structure of the 
lens perceived, if the light fidls in a proper direction. 

As the obliteration of micrometric lines in the experiment could 
only be produced by the retina retaining and combining a series of 
impressions whose joint duration is forty billionths of a second, a less 
time is sufficient for the production of each. If the number of views 
of lines presented to the eye be limited to ten, then four billionths of 
a second would be sufficient for each. Nor is this astonishing ; for, 
aranming the undulatory theory to be true, in this short peri^ two 
and a half millions of waves of light reach the eye. 



/ 



i 



REPORT ON THE PROGRESS OF PHYSIOLOGY. 225- 

Seksibilitt op the Eye fob Colours. — ^Lamansky ( Von Gr^fe's 
Archiv. xvn. l 123, 4) finds that the sensitiveness of the eye is 
greatest for green, yellow and blue, and least for red. This agrees 
with previous experiments, and with the known facts that with faint 
illumination, blue is better seen than red, that the peripheric portions 
of the retina are entirely insensible to red, and that in commencing 
atrophy of the optic nerve, red colour-blindness first appears. 

The time requisite for the perception of red is three times as 
much as that requisite for blue. The sensibility for white light is 
intermediate between that for green, yellow and blue on the one 
hand, and for violet, orange and red on the other. 

Spectra op Motion. — Dvorak {Wien. Acad-. SUzber. 2 Abtbeil. 
lxi. 257 — 262) seeks to explain the subjective sensations of motion 
whicb are observed when the eye has been fixed long on a moving 
object. Helmholtz explains them by unconscious movements of the eye, 
and brings them under vertiginous phenomena. Dvorak mentions 
several facts which, in his opinion, shew that the spectra are really 
local phenomena, and of the same nature as light and colour spectra. 

On Retina Currents. — Holmgren (UpscUa LclkareforeningB 
FordhamUingar, 1871, abstract in CeTUmmatt, Nos. 27, 28, 1871) 
finds that the whole theory of currents between the optic nerve and 
retina has been based on the false conception that the retina is the 
mere transverse section of the optic nerve. The retina, instead of 
being the simple transverse section, is a complicated structure, and is 
at the same time the longitudinal and transverse section. In the 
unstimulated eye each point on the posterior surface of the eyeball 
is negative to any point ot the cornea. The strongest current is pro- 
duced when the centre of the cornea is joined with any point in a 
circle half way between the optic disc and the ora serrata. This 
circle he calls the electro- motor equator of the eye. In the cornea 
itself and in the retina, there are currents according as the points are 
sjrmmetrical or unsymmetrical to the centre or equator. The centre 
of the cornea is always positive to any other part, and the electro- 
motor equator. negative to any point nearer the optic disc or ora 
serrata. 

When the i*etina is stimulated by light an oscillation takes 
place in all vertebrates except fishes. In the frog the oscillation is 
positive. In all others it is negative, but {»ositive when the light is 
withdrawn. The oscillation which occuiti on contraction of the 
pupil is not to be confounded with the retinal currents, for on con- 
traction of the pupil, i.e. the falling of light on the retina, the oscil- 
lation is positive, on dilatation, Le. disappearance of light, the oscil- 
lation is negative, which is just the opposite of the retinal currents. 
Experiments with light of various colours proved the inactivity of 
the ultra-red rays ; the strongest action is caused by the rays of the 
middle of the spectrum. The ultra-violet rays are still active. There 
is, therefore, an intimate relation between stimulation of the retina 
and oscillation of the current, the latter being *^ the objective signal 
of the subjective sensation.'' 

VOL. VI. 15 



226 BB BRUNTOM AND DR FEKBIER. 

Movements of the Eyeball. — Berlin {CwUraXblaU^ No. 35, 
1871) has made experiments in relation to certain movements of the 
eyeball observed by J. J. Muller and Donders on voluntarily opening 
or closing the eyelids. He determines the axis of rotation by a new 
method. The axis (drehpunkt) is constant for each horizontal plane, 
but on elevation of the plane of vision it lies more posteriorly, on 
lowering, on the other hand, more forwards. In voluntary forcible 
opening of the eye the eyeball projects, and on closure it again 
retreats. With the movement of the eyeball forwards there occurs 
also a movement downwards with a minimum shifting inwards. The 
synergic contraction of both oblique muscles and the levator palpebr® 
Buperioris is sufficient to account for all these movements. By move- 
ments of vision in a vertical plane the eyeball undergoes analogous 
changes of position. When the eye is directed from below upwards 
the globe projects from the orbit downwards and forwards, and vice 
versd when the eye is directed from above downwards. Further, it 
was found that in almost all horizontal and vertical visual move- 
ments there occur movements of the eyeball out of the visual plane, 
of such a nature that whenever the oblique and the lev. palp. sup. 
take part in a movement, the eyeball changes its position iu the line 
of traction of these muscles. 



Hearing* 

Movements of the Ear. — Mach and Eessel {CerUralblatt, No. 
38, 1871) by means of a new method have observed that the mem- 
brane of the fenestra rotunda bulges outwards when the stapes 
swings inwards, that contraction of the internal muscles alters the < 

excursions, and axes of rotation, of the ossicles, and that the excur- 
sions of the posterior segment of the membrana tympani in the living 
ear are much greater than those of the other parts of the mem* 
brana 

HvPESiESTiiESLA AcusTicA. — Folitzer {Arch, far OhrenheUk v. 
206 — 12) communicates an interesting case bearing on the physiology 
of the acoustic nerve. A woman had become deaf after an attack of 
typhus. Neither noises nor musical notes of ordinary intensity were 
h^urd, nor even the vibrations of a tuning-fork communicated through 
the bones of the head. When the notes of a harmonium were con- 
veyed to the ear by an ear-trumpet, it was found that thera was no 
reaction ^hen simple notes were sounded, but when the thirds of the 
higher notes of the scale were sounded tdie patient complained of an 
indescribable painful sensation in the ear and brain* In order to 
explain this, it must be supposed either that this is a reflex pheno- 
menon, or that the auditoty nerve, though chiefly a nerve of special 
sense, contains another class of fibres which are likewise capable of 
being stimulated by sounds, but instead of causing a sensation of 
sound convey a common sensation. Why these should be acted oa 
only by combined notes remains unexplained. 



BEPORT OX THE PROGRESS OF PHYSIOLOGY. 227 



Taate. 

Kerves op Taste. — LuHsana (Gazz, Med, Ital. Prov. Venet, xiii., 
CetUrcdblcUt, No. 15, 1871) brings forward a case from the clinique 
of Prof. Venzetti, which confirms the view that the lingiialis is the 
only nerve of taste lor the anterior part of tlie tongue. Most phy- 
siologists are agreed, that the fibres subservient to the sense of tuste 
are not contained in the lingnalis at its origin from the third divi- 
sion of the trigeminus. L. records two cases in which there was com- 
plete paralysis of all the three divisions of the- trigeminus. Though 
l^e common sensibility of the tongue was entirely loist, the sense of 
taste was intact. On the other hand, there are numerous cases of 
paralysis of the facial nerve in which the sense of taste was lost in 
the fore part of the tongue on the paralysed side. Lussana brings 
forward three new cases of this kind. Hence it appears that the 
nerves of taste to the anterior part of the tongue are derived from 
the facial and join the lii^ialis through the chorda tympani. SchifT, 
on the other hand, has stated, that the special nerves of taste to this 
part of the tongue, though not contained in the lingnalis at its origin, 
still are derived from the trigeminns. According to him they go 
through the second division of the fifbh to the spheno-palatine gan- 
glion, thence by the vidian nerve to the ganglion geniculatum of the* 
fitcial, and so through the chorda to the lingnalis. Lussana's cases 
overthrow this view. 

CirculcUton. 

Gakolia op the Heart. — Schmiedeberg {Lvdwig's Arbeiten, 
1870, p. 41) has found that the vagus contains fibres which have a 
quickening effect, as well as others which slow the action of the 
heart. The latter are paralyzed by nicotin, the former ai^ not. 
The inhibitory apparatus of the heart is not identical with the ter- 
minal filaments of the vagus in the heart, or with the motor ganglia, 
but is probably of a ganglionic nature and connected to the vagus by 
an intermediate structure. Nicotin paralyzes the filaments, or this 
intermediate structure, but not the inhibitory apparatus. Atropfn 
paralyzes the inhibitory apparatus itself The fibres which produce 
quickening probably act like the inliibitory ones through the medium 
of a ganglionic apparatus which is not identical with the motor 
ganglia of the heart. The stoppage of the heart which follows a 
cut, or the application of a ligature between the auriculo-ventricular 
groove and the venous sinux, is due to the motor-ganglia not being 
present to the same extent in all parts of the heart, and not to 
irritation of the inhibitory apparatus. 

Action op the Fascia op the Thigh on the Circulation. — • 
Braune {Ber. der adcJis. Gesell. d, Wiss. 1870, p. 261, AudCerUralbUut, 
p. 399) finds that when the leg of a dead body is rotated outwards 
and moved backwards, the femond vein becomes empty, and a nega-i 
tive pressure of ^ to 1 centimetre water is product in it. On 
restoring th« limb to its former position, or moving it forwards* th^ 



228 DR RRUNTON AND DR.FERRIER. 

vein becomes quite full. Repetition of these inovements, sncb as 
occurs to a certain degree in walking, sucks up the blood from the 
leg and presses it onwards to the body, the valves of the veins pre- 
venting it from flowing backwards. 

Absorption op Lymph by Tendons and FASCiiE. — In a research 
conducted under Lud wig's direction, Genersich {Ludwig's ArbeUeUy 
1870, p. 53) found that the lymphatics discovered by Ludwig and 
Schweigger-Seidel in fascise could be made to suck up a solution of 
alkanet in turpentine or ether. Thus a natural injection of them could 
be obtained. Thitf was done by caTefuUy freeiug that side of an aponeu- 
rosis which was directed towards a muHcle from connective tissue, 
stretching it tightly over a funnel, pouring upon it the alkanet solu- 
tion, and then causing it to move up and down by sucking the air 
from the funnel. The lymphatics on the side next the muscle run in 
parallel lines^ but on the side next the skin they form a uet-work. 
They do not absorb granular fluids, while the centrum tendineum of 
the diaphragm does so readily. In oi'der to discover what influence 
the contraction of muscles exerts on the sucking action of the aponeu- 
rosis, artificial circulation of blood was kept up in the hinder 
extremities of a dead dog which had been cut in two opposite the 7th 
dorsal vertebra, and the amount of lymph which issued from a cannula 
thoracic duct when the muscles were at rest^ was compared with 
that which flowed when they were made to contract. The blood 
flowed into a cannula tied in the aorta, and out through one in the 
vena cava. At firut^ a slight pressure of blood was sufficient to keep 
up a brisk circulation, but after about two houi-s the flow became 
slow, and although it could be quickened again by i*aising the pres- 
sure, the acceleration was only temporary. This slowing of the 
circulation was due to contraction of the arteries. The contraction 
began in the arteries of the skin, and extended to those of the 
muscles. It was increased by the circulation of perfectly fresh 
blood, but was diminished, and the current quickened, by stopping 
the circulation altogether for a while. The acceleration, however, 
was merely temporary, and soon became slow again. Strong electric 
irritation of the arteries caused them to dilate. He does not know 
what the cause of the contraction is, but it is not due to rigor mortis, 
as the vessels dilated when it came on. The muscular contractility 
becomes much diminished when the circulation gets slow, but this 
diminution is partly independent of the slowing of the circulation. 
In it^ passage through the vessels the blood lost oxygen and became 
venous, the amount of albumen was much increased, and the sugar 
(substance reducing copper) which it contained was very much 
diminished. Other conditions beiug the same, the amount of lymph 
obtained when the muscles were caused to contract by electric irrita- 
tion was much greater than when they were at rest, especially at the 
commencement of the experiment. This, however, is not so much 
due to the contraction of muscles pressing the lymph out of the 
lymphatics, as to the alternate tension and relaxation of the aponeu- 
roses, since passive movements of the limbs produce a considerably 



V 



KEPORT ON THE PB0GBES8 OF PHYSIOLOGY. 229 

gresAer flow of lymph than even the muBcuUr oontractions. When 
alkanet solution was injected between the aponeuroses and muscle 
it could be sucked up and flowed out at the thoracic duct of the 
limb hj making passive movements, but if injected between the 
skin and the aponeurosis, none was absorbed. Although a large part 
of the lymph obtained is probably preformed, and merely sucked up 
from the tissues, yet a certain amount seems idso to be formed during 
artificial circulation. 

Absorption without Circulation. — Goltz {Pfluger^s Archiv. 
IV. p. 147) communicates a preliminary notice of certain experi- 
ments on the absorption, and conveyance through the system, of 
poisons after cessation of the circulation. His experiments consisted 
in injectiDg strychnia into frogs after tying the heart and vessels. 
These proved that even after complete cessation of the circulation 
the poison is carried to every part of the body, as shewn by the 
fact that it produces its local effects on the spinal end wherever 
injected, and likewise by the fact that other frogs fed with any 
portion of the poisoned frog exhibited symptoms of strychnia poi- 
soning. 

The author reserves for further experiments, and a future notice, 
his views r^arding the forces concerned in the conveyance of the 
poison throughout the system. 

Yaso-uotor Nerves of Musoular Arteries. — Ludwig and Hafiz 
{jAidwig'8 Arbeitetiy 1870, p. 93) continued the researches on the cir- 
culation of blood in muscles, which Sczelkow, A. Schmidt, Sadler 
and Genersich had already made under Lud wig's direction. Their 
experiments were chiefly performed on dogs and rabbits, and they 

% recommend young animals for this purpose, as their vaso-motor 

system is moi*e sensitive than that of old ones. In order to ascer- 
tain whether the vaso-motor nerves of the arteries which go to 
muscles agree in their pi'operties with those of the vessels which 
supply the skin and viscera, all the vaso-niotor nerves* of the body 
were irritated at once, and the effect of this irritation on the arte- 
ries of the muscles compared with that on the vessels of the skin 
and viscera. The vaso-motor nerves were irritated by means of 
two electrodes inserted into the spinal cord, one opposite to the 
atlas, and a second at the under edge of the axis. The cord was 
previously divided between the occiput and atlas, and respiration 
kept up artificially. In order to eliminate the disturbing efiect 
which the contraction of the muscles produced by imtation of the 
cord would have had on the circulation in them, the animals were 
generally poisoned by an amount of curare just sufficient to paralyze 
the motor nerves of mascles, but not enough to aflect the vaso-motor 
system. The changes in the calibre of the arteries consequent on 
the irritation of vaso-motor nerves was noted in four ways. (1) By 
observing them with the naked eye or a lens. (2) By measuring 

^ the amoimt of blood which flowed out of the muscle when cut across, 

the larger arteries in it being ligatured previous to the measurement. 
(3) By the rapidity of the current i^is measured with the "strom-uhr." 



S30 . DB BRUNTON AND DR FERRIES. 

(4) By the qaicknees ^ith which the hlocMl pressure fell when the 
heart vms stopped by irritating the vagus. 

The experiments made iu these ways show (1 ) that when the cord 
is irritated tetanically the arteries of mnscltis contract, but to a 
much ieiss extent than those of the ^kiti and visceiti, so that the 
diminution in their calibre is not apparent to the eye. The other 
arteries contract every time the cord is irritated, but tho^e of the 
muscles do not. (2) The vaso-motor nerves of muscular arteries are 
easily exhausted, and when coutraction does occur during irritation 
of the cord, it only lasts a short time, and relaxation occurs while 
the luQ)en of the dermal and visceral arCeries is still continuing to 
dimiuish and the blood-pressure to rise. In consequence of thiSy 
the mubcles bleed profusely when divided dining irritation of the cord 
instead of scantily as tht-y do when the blood-presHure is at its 
normal height. (3) During irritation of the cord the current in 
muscular aiteries becomes n:ore rapid, and if the heai*t be stopped 
the bluod-pressure falls nearly as quickly as before, although the 
How of blood through the dermal and visceral arteries is much 
diminished. (4) The coats of muscular arteries exhibit active contrac- 
tions which are independent of the vaso-motor centre. They are distin- 
guished from those produced by irritation of the cord by not extend- 
ing over the whole length of the artery, but only affecting limited 
parts, so that the vessel sometimes looks like a string of beads. The 
periods at which they occur are quite independent of the irritation of 
the cord, and they persist during it notwithstanding the rise of pre»- 
sure which then takes place. (5) The biceps femoris i-eceives its motor 
nerves from the sacral plexus, but its vaso-motor nei'ves oome from 
the lumbar plexus through the crural nerve, and accompany branches 
of the crural ai*tery to the muscle. The meter and vaso-motor nerven 
can therefore be irritated 8e])arately. (6) In contradistinction to 
their nerves, the muscular fibres of the arterial eoats of muscular 
arteries are very irritable, and to their properties the alterations in 
the circulation are in all probability chieiiy due. They remain mode- 
rately contracted during the relaxation of the muscle provided that 
the muscular substance retains its normal composition. They con- 
tract strongly when exposed for a longer or shorter time to a high 
blood-pressure, probltbly fi*om the extension they undergo acting as 
an irritant iipou them. They dilate when the muscle contracts, or 
when it stops contracting, and also when the circulation in the muscle 
has been stopped for some time. In these cases alteration in the 
gases of the blood is not improbably the cause of the dilatation. 

Inhibition and Excitement in the Vaso-Motor Centres. — 
Cyon {Bfdl, de VAcad, imp, dea Sciences de St. Petershofurgy vii. Dec 
1870) considers that when a sensory nerve is irritated two kinds of 
reflex vano-motor action are produced, one general and the other local. 
The general consists in contraction of all the vessels of the body, 
the local m dilatation of the vessels of the part supplied by the 
■ensory nerve. From the former interfering slightly with the latter, 
a transient contraction pr e fe e d es the permanent local dilatation of the 



REPOItT ON THE PROORBSS OF PHYSIOLOOY. 231 

TaBsels. When the animal is narcotized, or the cerebral hemispheres 
removed, irritation of a sensory nerve produces dilatation instead 
of contraction of all the vessels in the body, and the local dilata- 
tion then occurs immediately, instead of being preceded by con- 
traction. When the whole cerebrum is extirpated, and only the 
cerebellum and medulla remain, irritation of sensory nerves pro- 
duces no reflex action whatever on the vessels. He thinks that 
every irritation of a sensory nerve when conveyed directly to the 
vaso-motor centre has an inhibitory action on it and produces reflex 
pai-alysis of the vessels, but when conveyed first to the brain, and 
thence to the vaso-motor centre, it has an exciting action, and pro- 
duces increased contraction of the vessels. It is alwavs conducted 
directly to the vaso*motor centre of the part which it supplies, and 
thus invariably produces local vaso-motor paralysis. 

Position op the Vaso-Motor Centre. — Lud wig and Owsjannikow 
(Ber, der sacks. Gesdl, der Wiss. 1871, p. 135, and CerUralblaU, p. 
483) find that the vaso-motor centre lies in the medulla oblongata 
from I millimetre under the corpora x^uadrigemina to 4 — 5 milli- 
metres above the calamus scriptorius, and to one side of the middle 
line. They determined this by cutting holes with a trephine 
through the skull of curarized rabbits or cats, on each side of the 
middle line, and then dividing the brain sabstance from the middle 
line outwards by means of small knives. Sections through the cor- 
pora quadrigemina produced a trannient rise of blood-pressure: be- 
tween the corpora quadrigemina and medulla oblongata a slighter 
rise. The rise was often accompanied by fieculiar oscillations. 
Sections in the medulla oblongata produced a fall in the blood-pres- 
sura The lower the section, the greater was the fall. The upper 
boundary of the district in which a section caused sinking was 
1 millimietre below the corpora quadrigemina, the lower border 4 — 5 
millimetres above the calumus scriptorius. A section made at the 
upper boundary had no effect on the action of the depressor, or on 
the rise of pressure after irritation of a sensory nerve; a section 
between the upper and lower boundary diminished these actions, 
and one at the lower boundary completely destroyed them. The vaso- 
motor centre^ therefore, lies in this district, and to one side of the 
middle line, as sections made in the middle line had no effect. The 
cerebellum has no connection with the blood -pressure. Sections 
above the medulla, or irritation of the sciatic nerve, always quickened 
the heart. Chloral produces a great sinking of the blood-pressure. 
Large doses of it destroy the power of sensory irritations to luise 
the blood-pressure, but small ones do not. The temperature falls 
after its adiministiution independently of the blood-pressure. 

l2ffFLUENC?E OP THE NeRVGUS StSTEM ON THE CIRCULATION AHD 

TpcPERATURE. — Riegel {Pfliiger's Arch, iv. 350 — 428) contributes the 
results of numerous experiments made by himself in reference to 
this subject, and combines with them a long critique of the nume- 
rous researches which have been made of late years in the same 
directi(m. He corroborates the ooeurrenoe of rhythmical eontractioui 



S32 DB BRUXTON AND DB FJSRBIER* 

• 

•in the Yessels as described by Schiff in the ear of the rabbit, Wharton 
Jones in the veins of the bat's wing, Saviotti in the web of the frog^ 
and describes them also in the mesenteiy of the frog and in the 
arteria saphena of the rabbit, where they are to be seen with great 
distinctness. (The occurrence of these rhythmical movements in the 
vessels of the rabbit generally had, however, been previously observed 
by Ludwig and Brunton. See Lud wig's Arbetien, 1869.) These 
movements are under the influence of the vaso-motor centre. Irritsr 
tion of the vaso-motor centre causes contraction, especially of those 
vessels which most distinctly manifept rhythmical contractions. The 
general eifect of such irritation, if it only affected limited vascular 
provinces, would be to aid the propulsion of the blood onwards into 
other regions, even though the circulation as a whole would be re- 
tarded. 

Riegel does not observe an invariable fall of temperature on irrita- 
tion of sensory nerves, and does not regard Heidenhain's explanation 
(see this Journal^ May, 1871) as satisfactory. 

Bapiditt op Circulatiok dt the Veins. — ^Cyon and Steinmann 
{BuU, cle UAcad, des Sciences de St Peter shourg^ xvi. 266) found that 
the amount of blood which flowed in a given time through the 
jugular, or cruiul veins of dogs narcotized with opium or chloral 
was nearly the same as that which flowed through the corresponding 
arteries. The velocity of the current in veins is therefore much 
greater than that calculated by Yolkmann. The rapidity of the 
current in the veins undergoes variations similar to that in the 
arteries, so that it is much diminished in the jugular when the 
carotid of the same side is compressed, but is increased by compression 
of the carotid on the opposite side. Division of the spinal cord 
lowers the blood-pressure, and thus lessens the rapidity of the 
current. Irritation of the cord generally raises the blood-pressure 
in dogs only to a slight extent, and as it causes contraction of the 
arterioles, it generally lessens the rapidity of the current. Some- 
times division of the cord is followed by a slightly increased ra- 
pidity of the current. Irritation of a sensory nerve produces general 
'contraction of the arterioles, and temporary contraction succeeded 
by dilatation in the part supplied by the nerve. When the tibial 
nerve was irritated the rapidity of the current in the crural vein 
was increased on one occasion so much that Oyon thought it due to 
local dilatation of the arteries occurring at the same time as a 
general increase ot blood-pressure. At other times the rapidity 
became greater as the blood-pressure sank, apparently from the con- 
traction of the arterioles, which had slowed the current in spite of 
the increased blood-pressure having given way to dilatation. If no 
rise of blood-pressure occurred, the stream in the veins was slowed 
by the contraction of the arterioles. Stoppage of artificial respira- 
tion in curarized dogs increased the rapidity of the current, and it 
only began to diminish when they were becoming asphyxiated. If 
the trachea of dogs which were not narcotized was closed when their 
Jungs were distended with air, the rapidity of the current in the 



REPOitT ON THE PROGRESS OF PHYSIOLOGY. 233 

reitis beciiliie rapidly diminished in consequence of venons con- 
gestion. 

Cattse op Vascular Murmitrs. — ^Nolet (Heynsius's Onderzoekingen^ 
II. p. 135, and UenU and Meisan&t^s Jahreaberickt, 277) finds that a mur- 
mur can occur in a tube of equal diameter throughout, if the stream 
passing through it has a sufficient velocity. The velocity necessary 
is greater than that found by Weber and Thamm. The narrower 
the tube, and the smoother the walls, the greater is the velocity 
necessary. The intensity of the murmur was equally great over the 
whole of a tube 18 inches long or more. A constriction in the tube 
causes a murmur both behind and in front of it, and the nan'ower 
the constriction, the less is the velocity of stream required. The 
murmur in front is weaker than the one behind, and a greater 
velocity is required to produce it. If the constriction was long 
enough, and of proper character, no murmur could be heard in its 
middle. A dilatation of the tube produced a murmur at the point 
where it began, and where it ended. A greater rapidity was neces- 
saiy to produce a murmur at its end than at its beginning. The 
wider the dilatation the greater the rapidity required, and if it be 
-very wide the velocity must be greater than that which is necessary 
in a tube of equal diameter throughout. Where an aneurism is 
very wide no murmur may be produced. The murmurs are due to 
vibrations in the fluid produced by vortices and strengthened by the 
walls of the tube acting as resonators. These vortices he rendered 
visible by means of powdered amber, and they corresponded in size to 
the intensity of the murmura. As the rapidity of the circulation in 
arteries at a distance from the heart is less than in those near it, 
roughness of the walls does not so easily cause a murmur in them. 

CoAorLATiON OF YdrtAL Blood. — Boll (Du JBoti* and Reichert^s 
Arch, 1870, p. 718) finds that the blood in the foetal chick before 
the 12th or 14th day does not coagulate, but merely deposits a sedi- 
ment of blood-corpuscles. At the 13th or 14th day it forms small and 
scanty coagula which are red from enclosed corpuscles. On the 16th 
or 17th day it begins to coagulate completely, but not so firmly as 
that of a developed bird. The non-coagulation in the first stage is 
due to want of fibrinogen. Fibrinoplastic substance is present, and 
when a few drops of blood from a chick about the 8th day are added 
to pericardial fluid, a jelly-like coagulum is at once formed. Hiemo- 
globin can be distinguished by the spectroscope in the blood on the 
3rd day of incubation. 

RespmUicn* 

Inplukitce op the Medulla on Respiration. — Schiff (i^itgrer** 
Archiv, 225 — 34) by further experiments confirms his statements which 
had been questioned by Brown-Sequard, that, at the level of the first 
cervical nerve the lateral columns of the medulla regulate the respi- 
ratory movements of the corresponding side. Section of the columns 
at this level causes entire cessation of the respiratory movements on 
.the same side, except in so far as that passive movements . may still 



234 DB BRUNTON AND DB FERRI£R. 

be observed, due to the active tinovements of the other side or abdo- 
men. In a rabbit in which the lateral column on one side was divided 
the volume of air respired was reduced one third of the normal. 

Inflxjence op the Vagus on the Air-cells op the Luno. — SchifiT 
(F^iiger's 4rchiv. iv.) shows that the expulsion of aii* from the lungs 
which results from stimulation of the vsgi immediatelj after death is 
partly due to contraction of the air-cells themselves. It is therefore 
probable that the vesicular emphysema observed in various parts of 
the lungs after section of the vagi in dogs and other animals is due to 
some extent to the paralysis of the air-cells which ensues. 

Post-moktem Appearances on Brain Lesions. — Fleischmann 
(Jahr,filr KhiderJieUk, iv.) confirms the facts observed by Brown- 
Sequard on lesions of various pi«rts of the brain (see Jounud ofAnM, 
wnd Fhy«., May, 1871). He found in cases of tubercle of the 
hemispheres, corpora quadrigemina, optic thalami and jions, apo- 
plectic extravasations on the pleui-e, pericardium and surface of the 
kidneys, and in one case patches of emphysema in both lungs. 

Extravasation in the Lungs after Diviskk^ of the Vagus. — 
Valentin (Zeitsclir^t /, Biologies vii., p. 160, note) thinks the scat- 
tered extravasations which appear in the lungs after division of the 
vagi are due to mechanical efforts, especially to the laboured respira- 
tion. In one case he found them in a quarter of an hour after the vagi 
had been cut. 

Influence op Change op Atmospheric Pressure on the Body. 
— ^G. V. Liebig {Arckiv. /. Klin, Med. viii. 445 — 60) seeks to detennine, 
with a view to the thei*apeutics of chronic lung disea.«^es, what influ- 
ence is exerted on the respimtion by increased or diminished atmo- 
spheric pressure. As is well known, respiration at high altitudes with 
low barometric pressure is exceedingly difficult This is due to the 
greater elastic recoil of the lung tissue from diminished atmospheric 
counterpressure. Hence the difficulty of inspiration and the disturb- 
ance of circulation. Under increased barometric pressure the pheno- 
mena are quite the reverse. The respiration is unusually easy and 
deep. The pulse also becomes slower. The influence of increased or 
diminished pressure is not of momentary duration, but persists for 
some time. Residence at high altitudes has therefore a tendency to 
develope the respiratory power and the strength generally. From 
the influence of diminished atmospheric pressure on the lungs the 
advantages of residence at high altitudes in cases of emphyseiya will 
be apparent. Em))hysematous lungs having lost their elasticity and 
offering resistance to the pulmonary circulation may regain their tone 
and respiraticm become easier. In all cases of diminished elasticity 
of tissue from any cause residence at high altitudes under low baro- 
metric pressure is advisable. 

Effect of Alterations in Atmospheric Pressure. — 'Bevt(Gofnpt. 
Rend, Lxxiii. p. 213) flnds that warm-blooded animals become con- 
vulsed and die rapidly if the atmospheric pressure be quickly reduced 
to 15 <Hr 18 cm. of mercury. After death the broachi are found filled 



7 



REPOBT ON THE FBOQKE9S OF F&J^HOLOQY. 235 

with bloody foarn^ and the blood ia dark on both sides of l^e heart. 
Death takes place equally quickly whether the bell-jar under which 
the animal is placed be closed completely or be traversed by a stream 
of air. If the f)ressare be dimininhed gradually, and a current of air 
be kept up, the animal will live for a considerable time under a very 
low prensure, but if the cuiTent be stopped they die of asphyxia. 
The composition of the air in which the animal dies varies with the 
pressui^e. Birds die at a pressure of 18 cm. of mercury, mammals at 
12 centimetres. 

Alterations in the Amount of Am in the Lungs during thk 
Movements of the Heart. — liandois {Berlin. Klin. Woch&nach. 1870, 
p. 9) connected a manometer with the trachea and found that air 
was not drawn in but driven put of the lungs during the systole of 
the heart. 

Ceradini {La meccanica del Cnuyre osmodei anruiU univeracM di 
medicina, see iv. Vol. 75, p. 587, and Henle's Jahresberichtf 1870, p. 
270) connected a manometer with the trachea, and saw that air was 
dj-awn into the lungs synchronously with the radial pulse. By using 
a sensitive registering instrument which he calls Hsemothorakograph, 
he found that during a small fraction of the duration of the systole, 
just at its beginning, the amount of arterial blood in the thorax re- 
mains unaltered, while venous blood still comes in, and therefore there 
is a slight expiration of air. Then arterial blood leaves the thorax 
more quickly than venous blood enters, causing a slight inspiration. 
As the systole ends there is a reflux of arterial blood towards the 
heart, and at the same time venoua blood streams into the thorax. 
This causes a slight expiratory movement, which, however, does not 
quite counterbalance the previous inspiration, and lastly, the air in 
the lungs returns with slight oscillations to its former volume. 

Influence of the Lachrymal Gland on Respiration. — Bergeon 
(Camptes Bendus, 1870, i., p. 88. Henle*s Jahresber. 1870, p. 162) 
thinks that one of the chief functions of the lachrymal gland is to 
keep the nasal mucous membrane and the inspired air moist, and 
thus preserve the mucous membrane of the respiratory passages from 
getting too dry. 

Temperature, 

Cooling of Warm-blooded Animals. — Horvarth {Centralblatt 
fiir Med, Wise, p. 531) has succeeded in cooling young dogs till the 
temperature in the rectum fell to 6 '6' 0. or even to i'S^ C, and again 
restoring them to their normal condition by means of warmth. On 
cooling rabbits to 7^ C. and a cat to 9*5^ C. no respiratory movements 
or cardiac pulsations could be observed, and the strongest electric 
currents applied to musdes or nerves had no effect. After they had 
been in this condition for an houi* he poui'ed warm water on them. 
The heart then began to pulsate and continued to do so for an hour, 
and electric currents applied directly to the muscles produced vigorous 
contractions, but the strongest currents applied to the nerves had no 



236 DB BBUNTOK AKD DB FEBBIEB, 

effect. Tliis affords an additional proof of the contractilitj of muscles 
independently of nerves. 

Influence op Heat on Animaus. — Bernard (Revue Scientifique, 
1871 — 72, pp. 133 and 182) investigates the action of temperature, by 
placing birds and rabbits in cages heated by a lamp or surrounded by 
a double metallic case containing a hot solution of sulphate of soda. 
When the animals are thus exposed to a dry heat of 150^ F., the re- 
spirations become quick and tumultuous, the temperature of the body 
rises, the heart beats quickly, and after a little stops suddenly if the 
temperature be high enough. The necessary temperature is sooner 
reached in birds than rabbits. It rose in birds to 122", in rabbits to 
115° F. Rigor moi'tis came quickly on, and both arteries and veins 
contained black blood. The blood from the vena cava contained 37*2 
per cent, carbonic acid, 3*4 nitrogen, and only 1 per cent, oxygen. 
The dark colour is due to rapid and extreme deoxygenation after 
death, as arterial blood collected immediately after death is of a 
scarlet colour. The rapid deoxygenation is due to the high tempera- 
ture, exactly the opposite condition being found in animals exposed to 
cold, the blood being then scarlet both in the arteries and veins. 
Blood drawn directly from an arteiy into a closed glass tube and gradu- 
ally warmed suddenly becomes dark, and between 140° and 158° F. it 
coagulates, and does not recover its bright colour when exposed to air. 
Involuntary muscles are very sensitive to temperature and variations, 
and produce contractions in them, but voluntary muscles are not. 
When animals are killed by warmth the irritability of the heart 
is entirely lost immediately after death. The stoppage of the heart 
by heat is, he thinks, due to coagulation of its myosin. From experi- 
ments on curarized frogs he finds that a temperature which destroys 
the irritability of muscles does not destroy that of motor nerves ; but 
sensory nerves, on the other hand, lose theirs at a lower temperature 
than muscle. 

Lauder Brunton (St Bctrtholomew^e Hasp. Reports, p. 216) tested 
the action of temperature on the action of the vagus, and on the rate 
of cardiac pulsations, by wrapping rabbits narcotized ^ith opium or 
chloral in cotton wool and laying them in a double case filled with 
hot water. The pulsations of the heart were registered on a revolving 
cylinder by one of Marey's levers, which was attached by a thread to 
a needle fixed in the ventricle, and the temperature taken by a ther- 
mometer kept constantly in the rectum. As the temperature of the 
animal rose the pulsations of the heart became mora and more rapid 
till they reached a maximum, their number then quickly diminished, 
they became irregular, and soon stopped completely, llie pulsations 
did not increase at the same rate throughout as the temperature rose, 
and the number of beats at any given temperature, as well as the 
amount of quickening for each degree of rise, vary in different 
animals. The temperature at which the heart stopped was between 
11 3° F. and 114° F. or even more, but it varied in different animals, 
and seemed to be much lower when a large dose of chloral had been 
given. 



REPORT ON THE PROGRESS OF PHTSIOLOGT. 237 

Division of the vagi before wanning the animal appeai'ed to }iave 
little or no effect on ^e quickeniog of the pulse by heat. In one 
case when they were not cut there were two maxima in the pulse-rate, 
one at 105*^F. and the other at 113^ As the temperature rose and the 
pulse became quicker the power of the vagus was diminished, so that 
the same irritation which at first had produced still-stand, on^ly caused 
slowing. When the same irritation was applied after the maximum 
was past and the heart had again become slow, just before death, 
still-stand was again produced. Irritation of the vagus immediately 
after death was followed by an undulating movement in the heart. 

BrBOULATiOK OP Tejtpebatdrb. — Eohrig and Zuntz {Pflugev^B 
Arch. IV. p. 57 — 90) uphold the doctrine of Liebermeister, that the 
production of heat in the body is dependent on the temperature of the 
surrounding medium. Cold within certain limits increases the tissue- 
change, and with it the production of heat. Heat has an opposite 
effect. They explain thus in the following manner. That by the 
cooling of the skin the cutaneous nerves are stimulated, and through 
them a reflex increase of the tissue metamorphosis is produced. From 
this it would appear probable that other stimulants of the skin might 
have a similar effect, and such in reality seems to be the effect of sea- 
and mineral -baths as diBtinct from fresh- water baths of the same 
temperature. 

The muscles are the chief seat of the heat-production, and this 
stands under a constant influence of the nervous system even without 
apparent muscular contraction. The reflex heat production by irri- 
tation of the cutaneous nerves is reduced to a minimum in animals 
paralysed by curara. 
/ The increase of temperature in the axilla which Liebermeister 

took for a proof of increased heat-pi*oduction when the skin was cooled, 
t))ey refer to the influence of the axillary vein on the thermometer. 
They found that movement of the aim without altering the position 
of the thermometer caused a rise in temperature in the axilla of from 
one-fifth to two-fifths of a degree. 

Action op Curare on Temperature. — Riegel {CentrcdblaU /tir 
Med. Wi88, 401) confirms the opinion of Bohrig and Zuntz, that the 
warmth of the body is chiefly produced in the muscles. He finds 
that when these are prevented from acting by poisoning with curare 
the temperature of the animal goes on constantly falling, although 
artificial respiration be carefully kept up, and the action of the heart 
and the functions of sensory and vaso-motor nerves remain unaffected 
by the poison. When the animal was previously rendered feverish by 
the injection of pus into its veins the temperature was not only 
reduced to the normal, but even below it. 

Means of lowering the Temperature. — Manassein {Pflilger^i 

I Archiv, iv. 283 — 301) writes on the influence which "rocking** 

exerts on the temperature of animals. He finds that by this means 

the temperature is lowered from 0*66 — 1 '2^ C It seems to be a phy* 

Biological effect brought about by the agency of the nervous system. 



238 DR BRUNTON AND DR FERRIER. 

iNFLtrENCB OF CooLiNO ON THE Pboduction OP Heat. — Wioternite 
(Wien, Med Jahrh, N.F., 1871, ii. p. 18) denies the trastvorthineas 
of Liebermeister's method of measuring the amount of heat given off 
from a body by the rise of temperature in the water of a bath in 
which the body is placed, an<jt shows that even after the water has 
been carefully stirred its different layers present considerable differ- 
ences in temperature. He does not attribute the rise of temperature 
which occurs in the axilla when the surface of the body is cooled 
to increased production of heat but to congestion in the periphery, 
as it can be produced by a tourniquet on the arm. No conclu- 
sion can be drawn from the temperature in one part of the body 
regarding that in another, as when cold was applied to the back in 
one case, the temperature rose in the axilla but fell in the rectum. 
The differences in temperature of different parts are due, he thinks, 
to reflex changes in the vaso-motor nerves. 

On Eegulation op Tehperaturb. — Senator, Virch. Arch, iavl 
p. 111. 

Liebermeister, Virch. Arcli. liii. p. 434, replies to the criticisms 
of T^internitz and Senator. 

Gildemeister {Diss, Basel, 1870, HenU cmd MeiMner^s Jahresber. 
1879, p. 208) finds that cooling the body causes increased production 
of Co, both in health and in fever. In fever, however, although the 
cold bath increases the combustion and production of heat in the 
body it withdraws more heat than is produced, and therefore cools 
the body and does good. 

Wertheim {W^m. Med. Wochmsch. 1870, pp. 297, 325, 353, 389, 
421) found that when the temperature of a dog was lowered by the 
application of ice the depth of respiration and amount of CO, expired 
was much increased. The CO, was increased both absolutely and 
also relatively to the amount of air respired. 

Influence op warm Sulphur Baths on Temperature. — Boettcher 
{Dorpat, Med Zeiiseh, 1870 i., 21, and CerUralblaU, 1871, p. 623) found 
that in sulphur baths from 34 to 35** 0. the temperature and axilla 
rose on an average 0-1** C; in the baths at 35® to 36*^0. it rose 
0-25" C, in those at 35 to 36-4'' C. it rose 0*34* C. Ab Kerner had 
found that in baths of ordinary water at 34® the temperature fell 
0-36* C, and in those about 36" C. it rose only 0121® C, it seems that 
the temperature in the axilla rises higher in sulphur-baths than in 
ordinary water. 

Perspiration in Fever. — Pudzinowitsch {CentralblaU, p. 211) 
draws the following conclusions from experiments made with Weyrich*s 
apparatus. The transpiration from the skin has no relation to the 
temperature. It may be diminished when the temperature is higher 
and increased when it is lower, as was seen in two cases of acute 
rheumatism, and one of pleuropneumonia. At a high temperature 
(39^ — 41^0.) it was diminished in two cases of remittent fever, and 
increased in two of typhus. In two cases of typhoid complicated with 
ague it increased or diminished in proportion as the temperature rose 
or felL 



f 



REPORT ON THE PROGRESS OF PHYSIOLOGY. 239 

Muscle. 

Theory op Muscular Power. — Luchsinger (Pfluger*s Archiv. 
IT. 201 — 205) does not confirm YoIckmann*s theory, that the exten- 
sion of an active moBcle by means of a weight which equals the abso- 
lute power of the muscle, always stands in a constant relation to the 
lifting-height of the unweighted muscle. He shews that this theory, 
if correct, would not harmonize with the commonly accepted Weber's 
theory of muscular activity, and that a muscle should be equally 
distensible by all degrees of stimulation, whereas it is known that 
the extensibility of a muscle is in proportion to the degree of stimu- 
lation. 

Elasticity op Muscle. — Fick {PJluger'a Arch, iv. 301 — 315) 
describes a form of myograph, in which the tension of the muscle is 
kept constant, and therefore adapted for determining how, during 
the state of activity, the length of the muscle varies, the tension 
remaining the same. Neither Helmholtz's nor Marey's myograph 
fulfils these conditions. The instrument in principle is a modification 
of Marey's, in which, however, the resistance to the muscle is not the 
elasticity of a spring whose tension varies during the movement of the 
lever, but a constant weight suspended on the lever. By a slight 
modification the instrument can be made to determine how, during 
the course of contraction, the tension of the muscle varies while the 
length remains the same. From a comparison of pendulum tracinga 
it is found, that when the irritation of a single momentary stimulus 
is allowed to proceed in the first instance, so that the muscle can 
contract while the tension remains constant, and next, so that with 
its length remaining the same the tension varies, the muscle in the 
latter case sooner attains its maximum of tension, than in the former 
case, its maximum contraction, i. e. its minimum length. Also, it 
appears that after momentary irritation the tense muscle becomes 
flaccid much sooner than the contracted muscle again i-etiims to its 
former length. The specific elasticity of the muscle diminishes in 
the course of contraction^ at first quickly, and then more slowly. 
The elasticity of the muscle is, therefore, in no respect a simple 
function of its natural length, but has dififerent values for each 
degree of contraction. 

. Diminution of Muscular Power durino Contraction. — Herr- 
mann (PflUger's ArcJUv, iv. 195 — 201) confirms Schwann's statement, 
that there is a diminution of muscular power during contraction. 
By graphic representation it is shewn that the curve of extension 
(Dehnung*s curve) of the active muscle has the form of a hyperbola, 
similar to that of the inactive muscle. This is a new confii'mation of 
Weber's theory of muscle. 

Muscular Tone — Sustschinsky (Centralblatty No. 34, 1871) in- 
vestigates the conditions of muscular tone in the hinder extremities 
of the frog and the influence of some poisons on the same. 

Muscular Rigidity. — Hermann and Walker {PJluger^a Archiv. 
IT. 182 — 195) investigate the phenomena of muscular rigidity. Ab 



240 * DR BRUNTON AND DR FERRIER, 

to tlie time of sbortening by spontaneous rigidity, it was found thai 
it does not proceed gradually after the sepai'ation of the muscle fiiom 
the body, but begins after a few hours, and advances at first with 
considerable rapidity, and afterwards moi-e slowly. They confirm 
the observation of Schmulewitsch, that in muscular rigidity induced 
by heat a distinct diminution of the volume of the muscle occurs. 
As to the absolute force of rigidity, they fouud that, as a rule, it is 
less than the power exerted by muscle on strong electrical stimula- 
tion, though sometimes fully equal or even greater. On a comparison 
of the amount of shortening of active and rigid muscles, it was found 
that light weights are raised higher by the ngid muscle, while heavy 
weights are raised higher by the living active muscle ; which proves 
that the ''natural form'* of the ri^id muscle is shorter and more 
extensible than the contracted living muscle. In regard to rigidity 
produced by a 8trong degree of cold, Hermann finds it depends on 
the rapidity of the freezing, whether a muscle will remain excitable 
after it is thawed or not. Contractions are caused by very rapid 
freezing, not if the freezing has been gradual. Neither the tem- 
perature to which the muscles are exposed after freezing, nor the 
duration of the frozen condition, nor the rapidity of thawing, have 
any influence on the rigidity. Muscles become rigid and acid even 
at such a lew temperature as 0^; and this is the case even with 
muscles which have previously been frozen and then exposed to this 
temperature. 

Electro-motor Properties of Muscles and Nerves. — ^Hermann 
(Pflilger's Arcfiiv. iv. 149 — 182) adds some new observations to his 
already published views regarding the nature of the electro-motive 
currents observed in muscles and nerves. He finds that in plants, 
as already stated by Buff, the transverse section is always negative 
to the longitudinal, and that artificial longitudinal sections are ne- 
gative to the uninjured surface. Experiments with heat and sub- 
stances destructive to the tissue, shew that the currents stand in 
close relation to the vital phenomena of the plant, and that their 
chief conditions are the presence of an injury to the yet living plant, 
just MS in muscle, the currents being genei-ated by the dissimilarity 
between living and dying protoplasm. Hermann has also made 
numerous observations as to the influence of temperature on the 
electro-motor properties of muscle. He had hoped by experiments 
on this point to bring to a crucial test his own theory, and the mole- 
cular hypothesiB of Dubois Keymond. Curiously enough, however^ 
the results obtained are explicable on either hypothesis. He found 
(1) between 0" and 30° the electro-motor power is greater at higher 
than at lower temperatures. Freezing and coagulation by heat destroy 
the properties utterly. (2) Cooled portions of muscle are negative, 
while warmed portions are positive to the other parts of the muscle. 
Freezing and heat-iigidity make the parts affected negative to the 
others. (3) Cooling or warming the transverse section has no ap- 
preciable influence on the electro-motor relation between this part 
and others. Freezing or heat-rigidity, however, increases its nega- 



I 



REPORT ON THE PROGRESS OF PHYSIOLOGY. 2il 

tivity, just like " fresheuing*' the section. These latter facts militate 
against Hermann's previously expressed view, that the negativity 
proceeds pari passu with the degree of molecular disturbance (Spal- 
tungsverschiedenheit), for then it should be expected that the colder 
muscular substance would be less negative, whereas exactly the 
opposite is the case. Hermann, ho wever,> still denies the pre-exist- 
ence of the muscular current. 

Laws of Muscular Exhaustioit. — From the experiments which 
Kronecker {Moruxtsher, der Akad, zu Berlin, 1870, p. 629) has per- 
formed with a highly complicated and most ingenious apparatus he 
deduces the following laws of muscular exhaustion. (1) If a 
muscle overloaded with a definite weight is irritated at equsJ inter- 
vals by equal (maximum) opening and closing induction shocks, the 
height of the contractions form an arithmetical series, in which the 
constant diiference depends only on the interval of time. If the 
muscle is loaded, instead of overloaded, the law only holds good 
for that height of contraction which is equal to the stretching 
caused by the weight. (2) When the intervals of time and the 
overload vary, the law is more general. The height of any con- 
traction with an overload p, and an irritation which follows the pre- 
vious one after an interval of time 0, is the same as if all the previous 
iiTitations had followed each other at an equal interval of time 0, and 
all the contractions had been made under the same weight. (3) 
Therefore the diminution in the height of contractions is independent 
of the weight, and only depends on the interval between two contrac- 
tions. (4) The changes of imtability disturb to a certain degree 
the expression of the previous laws. (5) Exhaustion can be partly 
removed by the injection of blood serum, dilute salt solution, to whicli 
a little permanganate of potash has been added, or even by pure salt 
solution. 

HLfiMOGLOBiN IN MuscLE. — ^Ray Lankester {Pfliiget^s Arohiv, 315 
— 20) shews that hsemoglobin occurs in the pharyngeal muscles of 
gasteropodous molluscs, such as Limnseus and Paludina, while no 
hiemoglobin exists in the blood. From its occuirence hei-e, and from 
the fact that it is found universally in the strongest and most active 
muscles (heart) of all vertebrates, and also in the muscles of the 
rectum and not in the rest of the idimentary canal, the author thinks 
there is some definite relation between hssmoglobin and muscular 
activity. 

Chest and FALSErrro Notes. — Mandl. {Wien. Acad. Sitzber. 
2 Abth. LXii. 264 — 70) describes the condition of the larynx in 
chest and falsetto notes. In chest notes the rima glottidis is open to 
its whole extent, and the vocal cords vibi'ate throughout their whole 
length. In the deepest notes the rima glottidis is elliptic, the widest 
part being about the middle of the ligamentous structures. The 
higher the notes, the neaver the processes vocales of the arytenoid 
cartilages approach each other, but the glottis respiratoria always 
remains open, and the vocal cords vibrate in their whole length and 
breadth. In falsetto notes, on the other hand, the glottis resjnratoria 

VOL. VI. 16 



242 DR BRUNTON AND DR FERRIER. 

is shut, and only the pars vocaiis of the vocal cords yibrates. In tbe 
deepest notes of this register the whole pars vocaiis vibrates ; in the 
higher, however, the false vocal cords apply themselves to the true 
vocal cords in front, behind, and at the sides, so that only the middle 
parts of the latter vibrate, and then only with their edges. The 
closure of the glottis respiratoria, which is necessary to falsetto, ia 
brought about by the action of the aiytenoid muscle, which moves 
the arytenoid caitilages on the facets of the cricoid from tlie under 
and anterior part to the upper and posterior. The crico-arytenoidei 
laterales alone can narrow the rima glottidis by swinging the aryte- 
noid cartilages on their vertical axis, but cannot cause complete 
closure. 

Digestion. 

Physiology op Vomiting. — Grimm (Pflilger'a Archiv. iv. 205 — 
209), considering the movements concerned in the act of vomiting to 
be merely abnormal respiratoiy movements, regulated, as Gianuzzi has 
supposed, by some centre in the medulla oblongata, has turned bis 
attention to the respiratory centre itself and experimented in how far 
the condition of apnoea might influence the act of vomiting. He 
found that when artificial respiration was vigorously maintained, 
the act of vomiting was very rudimentary and insufficient to empty 
the stomach, and also that on exhibition of tartar emetic artificial 
respiration is never able to cause entire cessation of the natural respi- 
ratory movements. These facts conGrm tbe idea that the acts of 
respiration and vomiting are innervated by the same centre and that 
the emetic causes a peculiar irritation of the respiratory centre. The 
exact nature of this irritation is a subject for further investigation. 

Movements of Intestine. — Sanders Ezn {Maandblad van de 
Genootachap ter hevordering van NcUuur Genees en-Hedkunde te Afnsier- 
dam, 1870-71, No. 6, and 8, and CentralbkUt^ p. 479) ties the oesophagus 
and rectum, and opens the abdominal cavity of animals whUe they are 
immersed in a 0'6 per cent, solution of Na. CI and thus avoids the irri- 
tation of the intestines, which would be caused by their exposure to 
air. In doing this the intestines are found to be at first perfectly 
motionless. Sometimes contractions of their longitudinal or circular 
fibres occur. The former are generally local, but their extent alter- 
nately increases and diminishes, and thus causes an undulating move- 
ment in the intestines. The latter often extend downwards along 
the intestines. Mechanical irritation of the intestines has little effect 
in producing movement. The amount of blood in the vessels changes 
without any apparent cause, but often remains constant for a long time. 
Dyspnoea causes the vessels to contract, but they again relax 'when 
the breathing becomes normal. The amount of blood in the vesaek 
has no influence on the movements of the intestina Compression of 
the aorta produces no movements in the intestine when at rest^ and 
may even dimini«h movements when the^ are present. The utems 
behaves in a similar way to the intestine. The violent movements of 
the intestines, which generally occur during suffocation, do not take 
place if the vagi are previoiuly divided in the neck. IrritatioD of 






I 






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REPORT ON THE PROGRESS OF PHYSIOLOGY. 243 



tbeir peripheral ends produces the movements. These occur chiefly in 
two places, the under end of the duodenum and the upper end of the 
ileum. Irritation of the right vagus chiefly causes movement in the 
duodenum, and irritation of the left movement in the ileum. The 
stomach sometimes remains quiet, but generally it expels part of its 
contents from time to time into the duodenum. In one case of diar- 
rhoea he saw antiperistaltic movements iu the colon, in all other 
parts the movements were peristaltic. The urinaiy bladder some- 
times presented movements even when empty. These are always 
peristaltic. The horns of the uterus present peristaltic and anti- 
peristaltic movements. 

Absorptiox op Fat. — S. V. Basch (If ten. Acad, Sitzber, lxii. 
2 Abth. 1870, 617—634) holds that the central lacteal of the villus 
stands in open communication with the lacteal passages which per- 
meate the villus, and is devoid of epithelium. The particles of fat 
permeate the base of the epithelium of the villi. There is, however, 
no direct communication between the epitheliun^i and the lacteals, there 
being only a close contiguity between the margin of the villus and the 
epithelial cells. The appearance of fat particles between the epithelial 
cells often observed after death 'is due to a sort of extravasation or 
regurgitation from the lacteals. 

Function op the Spleen. — Hosier (CerUralblcUt, No. 19, 1871), 
in a preliminary notice, gives the results of 30 cases of extirpation of 
the spleen, and draws the following conclusions : (1) The spleen is not 
necessary to the life of the animaL (2) After extirpation as well as 
after artificially caused atrophy of the spleen, its frinction is under- 
taken by other lymphatic organs, especially by the medulla of bones. 
Marked changes are seep in this after extirpation of the spleen. 
Hyperplasia of the lymphatic glands is not a constant phenomenon. 
J*? ^3j The vicarious action of the lymphatic glands is not complete at 

*^ first, as the blood is altered in character for the first few months after 

extirpation. The spleen has a direct influence on sanguinification, 
and is apparently concerned in the formation both of white and red 
corpuscles. (4) The spleen has np influence on the gastric or pan- 
creatic digestion. The great appetite which is said to follow extirpa- 
tion of the spleen is not a constant occurrence. 

Secretion op Urine. — A paper by C. Ustimovitsch (in I/udtoig^a 
Arheiten, 1870, p. 198) contains much interesting information on 
this subject. He shows that the secretion of urine is the product of 
two factors, one of which is the difference of pressure between the 
circulating blood and the excreted urine, and the other the amount of 
urinary material which the blood contains. Max Heimanu had found 
that the secretion of urine depends on the difference of pressure be- 
tween the blood in the renal arteries and the urine in the ureter, and 
that secretion might be arrested either by lessening the pressure of 
the blood or increasing that of the urine. Ustimovitsch shows that 
when the amount of urinary material in the blood is about the same, 
the secretion of urine goes on in proportion to the difference between 
the pressure in the blood-vessels and that in the ureters. No secre- 



244 DR BRUNTON AND DR FERRIER. 

tion takes place when the blood -pressure sinks below 40 mm. of 
mercury. He points out that the pressure of blood in the renal 
arteries is quite a different thing from that in the systemic arteries 
genei^allj. It must sink along with the pressure in the aorta, 
but the muscular coats of the renal arteries may contract and cause 
it to fall though the general tension in the aorta remains high. This 
contraction may be due either to the vaso-motor centre or to 
causes in the kidney itself. Section of the renal nerves separates 
the arteries from the vaso-motor centre, and therefore generally 
causes the contraction to relax and increases the amount of urine. 
Sometimes, however, local causes prevent the contraction from 
relaxing, even when the nerves are cut, and in those cases section 
of the nerves does not increase the secretion. Section of the 
splanchnic has the same action as division of the renal nerves, but it 
also produces diminution of the blood-pressure, and therefore does not 
increase the secretion so much. Section of the cord at the lower 
part of the neck stops the secretion of urine. If Na. CI or urea be 
then injected into the veins, secretion begins again notwithstanding 
the great diminution in blood-pressure which follows section of the 
cord. For a criticism of Eck hard's theory of secretory and inhibitory 
nerves, we must refer to the original. As regards the proportion of 
of water, urea and chlorine secreted in a given time, TJstimovitsch 
finds that when the blood-pressure sinks, all three are more scantily 
secreted. When the urine first secreted is rich in urea, its amount 
becomes diminished more quickly than that of the water, but if 
it is poor the water diminishes before the urea. Section of the 
renal nerves generally increases the urine, and in these cases the 
water increases much more quickly than the urea* When the resist- 
ance in the ureter is raised so that secretion becomes scanty, the urea 
diminishes much more quickly than the water. The same is the 
case when the urine which flows out without any resistance contains 
3 per cent, of urea. When the amount of Na. CI in the blood is 
augmented, the seci'etion of urine becomes increased ; the Na. CI rises 
quicker than the water, and with every additional gramme of Na. CI 
excreted a very constant weight of water (14 — 16 grm.) passes out. 
Increase of the amount of urea in the blood has an inconstant effect. 
Curare poisoning stops, or greatly lessens, the secretion of urine, and 
the chlorine and urea are more diminished than the water. When 
the renal nerves are cut, secretion again commences, but the 
urine remains watery and very poor in urea and Na. CI even though 
these be injected into the blood. This he ascribes to a specific 
action of curare on the urinary secretion. When artificial respi- 
i-ation was stopped, and the curarized animal became asphyxiated, 
the urine began to flow more rapidly. 

Participation op the Kidneys in the formation of Urea. — 
Rosenstein {Cen^cUblaU, No. 23, 1871, p. 353) investigates what 
part, if any, the kidneys play in the formation of urea. He considers 
all previous methods faulty which consist merely in analysis of the 
blood going to, and the blood returning from, the kidneys. His 



^ 



i^ 



REPORT ON THE PROGRESS OF l^HYSIOLOQY. 245 

method was to compare the amount of urea in the urine of a dog 
retaining both kidneys with the amount af%er extirpation of one 
of them. The result was, that as much, if not more, urea was 
excreted when only one kidney was present as when both existed. 
This might be attributed to complementary hypertrophy, but the 
results were observed long before any hypertrophy could have 
taken place. The conclusion is, that the kidneys have nothing to 
do with the formation of urea. 

Miscdlaneaus. 

Absorption op Insoluble Substances. — Auspitz (Wien, Med. 
Jaharb. N. F. 1871, iii.) gives the results of experiments on rabbits 
with regai*d to the absorption of starch suspended in water or oiL 
Starch injected into the veins reaches the lungs, where the greater 
part is retained, without producing serious inconvenience or inflam-> 
mation. A small portion of the granules reach the liver, spleen, 
and the vessels generally. Injected into the abdomen, or subcuta- 
neous cellular tissue, starch suspended in water, but particulai^ly if 
suspended in oil, reaches the circulation and the viscera. Oil injected 
Bubcutaneously diffuses itself in the meshes of the subcutaneous and 
muscular connective tissue, and reaches as far as the serous linings 
of the hollow cavities. Inflammatory symptoms were not observed 
in the organs, but often hemorrhagic extravasations. Wherever in- 
jected, starch was found in the thoracic duct After infriction of 
starch and oil into the skin of rabbits, immediately before or after its 
separation from the body, these substances were found in the oorium, 
the subcutaneous cellular tissue and the attached muscles, but not 
in the sebaceous or hair-follicles. The process of friction was 
carried on for a quarter of an hour. Fats gi-eatly favour absorption 
of substances. 

Electro-motor Fhexohena IK THE Skin op Frog. — Engelmann 
(Pfliige/s Archiv, iv. 321 — 32 4) has continued his researches on the 
contractile glands of the frog's skin (see Journal of Anout, and Phy$, 
May, 1871). The electro- motor properties of the frog's skin have 
long been known, but it struck Engelmann that possibly the currents 
were due to the contractile cells which form a special layer of the 
so-called epithelium of the glands. Numerous observations of the 
coincidence of variations in the electro-motor power, with changes in 
the appearance of the contractile cells, have convinced him that 
these cells are the only source of the cutaneous currents of the 
frog. These electro -motive propei'ties stand in intimate relation to 
secretion. He regards the process of secretion in these glands as 
nothing more than electrical osmosis. The direction of the current, 
though at first sight contrary to the course of secretion, is readily 
explained by the position of the cells, the current passing in their case, 
as in muscle, from the longitudinal to the transverse section. 

CoNTBACTiLiTY OF White Oorpuscles. — Hoppe-Seyler {Med. 
Chem. UrUersuch. p. 486) finds that white blood or lymph corpuscles 
contain glycogen so long as they exhibit movements, but when 
they become rigid they lose their glycogen and contain sugar. In 



246 DR BRUNTON AND DR FERRIER, 

the pus from abscesses and wounds no glycogen was found, and tlie 
occurrence of glycogen may, therefore, be taken as a means of distin- 
guishing lymph from pus corpuscle, although the latter spring from 
the former. 

Action op Sea Watkr on Fresh-Water Animals. — P. Bert 
{Comptes Rendus, lxxiii. 382 — 5, 464 — 7) investigates the cause of 
death when fi-esh- water animals are placed in sea water. Fresh- 
water animals placed in salt water of a temperature of 15*^ to 16"0. 
die after some minutes or hours, according to the species. Fresh- 
water animals may, to a certain extent, be acclimatised to salt water 
by gradual addition of the latter, but they invariably die when the 
proportion of salt water is more than one half Experiments with 
the individual constituents of sea water proved that the fatal effect 
is chiefly due to the chlorides. The cause of death is exosmotic 
dessication. In animals breathing by gills the immediate cause 
of death appears to be sudden arrest of the branchial circulation. 
The more delicate the structure of the gills the more rapid the effect. 

Action of Fresh and Salt Water on Animal Life. — In a 
series of experiments, F. Plateau {Mem. de la Soc, Roy, Belg. xxxvi. 
and Academy y 311) found that the fatal effects of sea water on fresh- 
water species of articulata, and of fresh water on marine species, 
are due to differences in chemical composition and not in density, as 
fresh-water species lived perfectly well in a solution of sugar of the 
density of sea water. The poisonous action of sea water is due to 
the chlorides of potassium, sodium, and magnesium, while the sul- 
phates of magnesium and calcium have little or no effect. The 
sulphates are hardly absorbed at all, but the chlorides are rapidly 
absorbed, and if the animals are placed in distilled water, are 
again given out. When marine articulata are placed in fresh water, 
they give off chloride of sodium, and the loss of this Plateau believes 
to be the cause of their death. 

Cause of Phosphorescence. — Panceri (see Academy, p. 401) 
thinks phosphorescence is due to the slow oxidation of fat In fishes 
oxygen penetrates the skin and oxidizes the fat below. It is increased 
by oxygen, but stopped by immersion in carbonic acid, fresh water, 
alcohol, or any substance not containing oxygen. Phosphorescence 
begins at death and stops when putrefaction begins. 

Experiments in Pangenesis. — In a series of experiments which 
he made in order to test the truth of Darwin's theory of Pangenesis, 
Gal ton {Froc. R. S. p. 394) injected the blood of other kinds into the 
veins of pure silver-grey rabbits and then bred from them. If gAi- 
nules were present in the blood, and the sexual elements consisted 
simply of organized groups of these granules, the offspring of these 
rabbits ought to have presented some of the characters of the other 
breed from which blood had been transfused into the parent animals, 
in addition to their own natural ones. His experiments consisted of, 
1st. Moderate transfusion of partially defibrinized blood. 2. A large 
transftision of wholly defibrinized blood. In both these cases a 
quantity of blood similar to that injected was previously drawn from 
the vein. 3. Establishing a system of cross cii*cuiation between the 



4 

f 



^ 



REPORT OF THE PROGRESS OF PHYSIOLOGY. 247 

carotid artery of a silver-grey and of a common rabbit The injec- 
tion of blood sometimes produced temporary sterility, but had no 
effect on the offspring, which exactly resembled their parents, except 
in one doubtful case. Cross circulation had no effect whatever. 
This shows that the reproductive granules are not independent 
residents in the blood, though it is possible that they may be tem- 
porarily present in it. 

Reproduction op Epithelium.— Heiberg {Strieker^ a Medicin, 
Jahrh, Heft i. p. 7) finds that when the epithelium is scraped off 
the surface of the cornea in the frog, fowl, or rat, it is reproduced 
only at the edges of the denuded part, and never from any isolated 
centre on its surface. The isolated centres from which reproduction 
of the epithelium of denuded skin takes place are, he thinks, gland- 
ducts lined with epithelium. He considers that the abraded epithe- 
lium is reproduced in the cornea by cells at the margin of the 
denuded spot sending out processes in which a transparent spot ap- 
pears. The processes then separate from the parent and develope 
into new cells, the spot becoming the nucleus, and then in their turn 
send off new cells which gradually cover the bare spot. 

Origin and Distribution of Sugar-forming Ferment. — Lepine 
(Lud wig's Arheiien^ 1871, p. 11 3) finds that starch is converted into 
sugar iu half-an-hour or an hour by pieces of the mucous membrane 
from the tongue or palate of the frog, or by the fluid which covers 
their surface. The fluid can be obtained by means of small pieces of 
satin-paper laid on the tongue. The mechanical irritation which 
these produce cause the tongue to become red and the fluid to be 
secreted so plentifully that it may be removed by means of a glass 
rod. The secretion can also be stimulated by electrical irritation of 
the hypoglossal and glossopharyngeal nerves and of the tongue itself. 
When it is examined microscopically, after its nerves have been 
irritated some time, numerous lymph-cells are found in the neigh- 
bourhood of the racemose glands which are contained in it, and 
which may be regarded as salivary glands. A sugar-forming ferment 
has been found by other authors in the mucous membrane of the 
stomach, intestine and bladder, liver, bile, kidney, brain and muscles; 
but he obtains it besides in dogs, rabbits, and frogs, from the spleen, 
lungs, testicles, tendons, serous membranes, cornea, and vitreous 
humour, from both the inner and outer surfaces of the frog's skin, 
the mucus of its ovary and that covering the slug. It does not 
exist in equal quantities in all organs, but the only place where it 
was absent was the crystalline lens. It diffuses through vegetable 
parchment. Human saliva, unlike that of the dog, does not become 
more powerful by standing. In accordance with Bemard^s statement 
he finds that a sugar-forming ferment is produced during the decom- 
position of albuminous substances, the crystalline lens included. 

On the Evolution of Ammonia from Putrefying Blood. — 
Exner {Wien. Acad. Bericht, 1870, B. lxu. 2 Abth. S.A.) has studied 
the influence of atmospheric air, and specially of oxygen, on the 
process of putrefaction of blood with particular reference to the obser- 
vations he had already made that access of oxygen favours the 



248 BB BRUNTON AND DR FERRIER. 

eTolation of ammonia. When atmospheric air was passed through 
blood contained in a wide-mouthed vessel, for two or three days, an 
intense evolution of ammonia took place and the blood-oolouring 
matter was set free. He shews that the evolution of the ammonia is 
not due to the discharge of the colouring matter. When equal 
portions of blood were placed under different degrees of pressure, 
the evolution of ammonia was greater under the higher pressure. 
In order to determine whether the organisms contained in the 
atmosphere played any part in the formation of ammonia, he passed 
into the blood air filtered through cotton wool or nitric acid, and 
found that the effect was in nowise modified. 

Experiments as to which gas the evolution of the ammonia was 
due, proved that pure nitrogen and also carbonic acid which Pasteur 
regards as necessary for fermentation, had no effect, and that pure 
oxygen had no more energetic effect than ordinary air. When blood 
was treated with hydrogen the evolution of ammonia was greater 
than under the action of nitrogen, but less than that caused by air. 
When a quantity of blood was sealed up from access of air, an 
evolution of ammonia was observed when the vessel was continually 
shaken. In a similar vessel allowed to remain perfectly at rest no 
ammonia could be detected. In all cases and in all fiuids experi- 
mented with, low organisms, such as bacteria, vibriones, &c, were found. 

Product op Biliary Pigment. — Stockvis (Maandblad der sectie 
voor Natuunvetenschappen, 1870, No. 305) has observed that the 
presence of bile-pigment in icteric urine can be recognised by its 
becoming of a brownish green colour, and presenting peculiar spectro- 
scopic absorption-bands after it has been treated with chloride of 
zinc and ammonia and exposed to the air. The same absorption-bands 
are presented by biliverdine after treatment with chloride of zinc and 
ammonia or by blifuscine (?) which has been exposed to the air for 
2 — 4 days. During the preparation of biliverdine from bilirubine 
through the action of chloride of zinc and alkalis, bilifuscine or some 
substance nearly allied to it probably occurs as an intermediate pro- 
duct. When it is treated with acids the substance appears to yield 
biliverdine, and a i>eculiar colouring matter soluble in water. This 
peculiar brownish green or green colouring matter is an oxidation 
product of bile-pigment and nearly allied to those which occur in 
Gmelin's reaction. 

Fluorescing Product op the Reduction op Blood-colouring 
Matter. — Stockvis {Maandhlad der sectie voor Natutianffetensehcippen, 
1871, No. 9) has obtained a peculiar rose-coloured, fluorescing sub- 
stance by boUing a solution of dried blood, hemoglobin, haematin or 
hsematoin in alcohol with hydrochloric acid and tin, diluting the 
product, and treating it with ether chloroform or amylic alcohol. 
He has not obtained it in a cryistalline condition. It presents peculiar 
spectroscopic absorption bands which are quite distinct from those of 
iron-file haematin though slightly resembling them. It resembles bile 
pigments in its fluorescence, in the altei'ations produced in its spec- 
trum by mineral acids, and in its solubility in either alcohol and 
amylic alcohol. He has not been able to oxidize or reduce it 



A NEW SCHEMA OP THE CIRCULATION. By Wm. 
RuTHERFOBD, M.D. F.RS.K Profe89or of Physiohgt^ K%ng^$ 
College^ London. 

MuoH may be learned regarding the cirenlation with the aid of 
Weber and Marey's Models. Weber's Model or Schema is a simple 
elastic tube (formerly a piece of intestine), at one part of which two 
membranous valves are fixed bo as to represent the valves of a cardiac 
ventricle; at another part of the tube a piece of glass tubing contain- 
ing a sponge is inserted ; the pores of the sponge representing the 
capillaries. By means of this model the student can readily under- 
stand why the blood moves in a certain direction, and he can also 
learn a good deal regarding the blood-pressure. The Schema con- 
structed by Marey is more elaborate and is capable of shewing a good 
deal more. It consists of a system of elastic tubes arranged so as to 
represent the Arterial System. There are no veins or capillaries, and as 
the apparatus is worked the water which is driven through the tubes 
pours out by the arterial orifices. The water pours from these into 
a glass funnel which transmits it into the auricle of the heart. Mano- 
meters and sphygmographs can be adapted to the arteries, and nume- 
rous important lessons can be learned from the employment of these. 
The apparatus, although ingenious, is nevertheless very incomplete, 
and inasmuch as it must always be kept in a vertical position, it is 
inconvenient. I have recently had constructed a Schema of the circu- 
lation, which serves to shew all that Weber and Marey*s models 
exhibit, and a great deal more. I therefore think it worUi while to 
direct the attention of physiologists to it, for I find it of the greatest 
service for teaching purposes; and it will, I think, enable us to arrive 
experimentelly at several important fiicts regarding the circulation. 
I am at present working at this subject, and feel warranted in making 
the above statements. 

The figure that I give below looks somewhat formidable. At 
first glance the reader will probably regard the Schema as labyrin- 
thic enough, but the complication will quickly disappear. I have 
represented it in connection with various instruments which may be 
used for studying blood-pressure and motion. 

Presuming that the reader has perused the description of the 
above figure, I will simply say that the tubes of the apparatus 
consist of vulcanised Indiarrubber. Those representing the arteries 
are thick and strong, and do not collapse when empty. Those repre- 
senting the veins have thinner walla^ a larger calibre, and, unlike 
the arteries, they collapse when empty. The ventride of the heart 
consists of an ordinaiy elastic pump. One with leaden balls does 
perfectly for illustrating all the facts regarding the blood-pres- 
sure, the abolition of the pulse-wave by the capillaries Ac, but for 
sphygmographic and sphygmosoopic tracings valves consisting of 
bftUs do not suffice, inasmuch as they give tracings very different 
from those which are obtained from human vascular system. Yalves 

VOL. VL 17 



252 PBOF. BUTHEBFOBD. A NEW SCHEMA, &C. 

(S) may be employed, and its tradngs recorded npon a revolying 
oylioder. Marey's Sphygmoacope S', or • Fick's Eeder-Manometer, 
may be connected with another, or it may be the same tabe, and the 
tracings produced by them may be recorded upon the same cylinder 
with the sphygmographic, and also the kymographio (manometer) 
tracings. Most Taluable comparisons can thus be instituted between 
the tracings produced by these various instruments. Lud¥rig's Hse- 
mometer or Stromuhr ^S'") can be attached to another, or it might be 
the same tube as the spnygmograph or manometer; and the volume of 
fluid that flows through the tube in a given time can thus be com- 
pared with the pressure, &o. Chauveau's HsBmodromometer or Tacho- 
meter may also be inserted, and the speed of the current compared 
with the pressure, &c 

The effect upon the current of causing it to pass through tubes of 
difierent calibres, tubes which have constrictions or dilatations, tubes 
which divide at acute or obtuse angles, &o. may be investigated. 
All these, and many other things, may be readily studied with the 
aid of this apparatus. The tubes K K^ K." can be opened, if other 
tubes are needed. When the apparatus connected with the tubes 1 
and r is not used, the tubes are clamped or tied and the piece of 
tubing added, taken away from a short stump of tubing (like that 
at K' and K'') left in connection with the proper vessel of the 
Schema. Junctions of tubes are made by placing a little piece of 
glass tubing within the adjoining orifices of the ends of the tubes, 
and these are of course tied firmly upon the glass tubes. 

This apparatus is valuable to the student as well as to the teacher, 
for by handling it for himself, the student can readily gain a know- 
ledge of the circulation^ which books can never enable him to obtain. 

The India-rubber part of the Schema has been cheaply and 
satisfactorily constructed for me by the North British India-Rubber 
Company, Canon-mills, Edinburgh, from whom it can at any time be 
obtained. 



/' 



/ 



/• 



journal ot ^natom^ anti ^i}^9iio\ogg. 



* 



* 



THE MUSCLES OF LEPIDOSIREN ANNECTENS, 
WITH THE CRANIAL NERVES. By Pbofessoe 
Humphry. (PL xii. Figs, 28 to 270 

The muscular system of Lepidosiren presents \ as might be 
expected, many interesting features of an intermediate condi- 
tion between that of the Fish and that of the XJrodelan. 

The great lateral muscle, as in those animals, is divided by 
a lateral septum passing, from before backwards, nearly mid- 
way between the dorsal and the ventral edges, into an upper or 
' dorsal* and a lower or * ventral' muscle. Each of these is also 
subdivided into myotomes by transverse fascial inscriptions or 
' intermuscular septa' which are more simply disposed than in 
most Fishes, forasmuch as they run from the lateral septum, in 
nearly straight lines or with a slight anterior curve, upwards 
and downwards, to the dorsal and ventral edges. Traced, from 
the external fascia, into the substance of the dorsal muscle the 
septa pass, in oblique planes, forwards and inwards to the dorsal 
spines and the fin-rays ; and those in the ventral muscle pass 
also obliquely forwards and inwards, to the ribs, haemal spines, 
and fin-rays, where these exist, or to the internal surface of the 
ventral cavity and there blend with the internal or transver- 
salis fascia. 

Fascial Investment. 

The EXTERNAL or SUBCUTANEOUS FASCIA Covering the late- 
ral muscle is not, as in Cryptobranch, loose, permitting move- 
ment of the skin upon the muscle, but, as in Fishes though not 

1 The Bpecimen of Lepidofliren Anneotena from which fheae notes were 
taken was nineteen inches in length. It was dissected chiefly hy ICr Anningson, 
partly hy myself. 

VOL. VI. 18 . 



254 PROFESSOR HUMPHRY. 

quite in the same degree, compact and closely united with the 
derma on the one side and with the intermuscular septa on the 
other. It is separated from the skin by a thin stratum of pig- 
mentary areolar tissue ; and its union with the intermuscular 
septa is still more close. Hence it is with difficulty dissected off 
as a continuous sheet. Forasmuch as all, or nearly all, the 
fibres of the lateral muscle terminate in and arise from the 
septa, they are, through the medium of the septa, brought into 
relation with one another, with the vertebral system on the one 
side, and with the dermal system on the other. This is an 
important point ; and the continuity of the parts or layers of 
the outer wall of an animal body thus established must be 
borne in mind in considering the relations of the muscles, and 
their subdivisions, to one another and to the dermal and verte- 
bral systems. 

Along each edge (upper and lower) of the animal, in the 
forepart, where the mesial fin is absent, the fascia of either 
side blends with that of the opposite side and is intimately 
united with the cutis externally. The median sheet resulting 
from the blending dips between the lateral muscles of the two 
sides. The upper one impinges upon and unites with the dorsal 
spines and other vertebral elements; and the lower blends 
with the haemal spines or with the fascia transversalis. 

In the region of the mesial fin (dorsal, caudal, subcaudal and 
anal, for they are continuous) the disposition is somewhat different 
At the base of the fin the fascia on each side divides into two 
laminae (Fig. 23 B). Of these one, superficial or subcutaneous, 
accompanies the derma upon the fin and is lost with it on the 
delicate homy fibres of the fin. The other lamina, retaining its 
connection with the lateral muscle, passes over the edge of that 
muscle, between it and the fin covered by its fin-muscle, and so 
reaches the neural or haemal spines and the fin-rays resting 
upon them. It here becomes blended with those spines as well 
as with the intermuscular septa which are here passing into 
the spines and fin-rays. Thus the fin with its muscles is 
enclosed in the interval between the superficial and deep fascial 
laminae of the two sides, in the same manner as the dorsal, 
ventral and lateral fat masses in Cryptobranch (p. 6). There is 
however this difference, that the fat masses in Cryptobranch 



K . 



•\ 



MUSCLES OF LEPIDOSIREX. 255 

occupy continuous channels which are not broken or subdivided 
by the transverse septa extending into or across them ; whereas 
the channel for the mesial fin of Lepidosiren is, in the line of 
each fin-ray, interrupted by the transverse septa of the lateral 
muscles of the two sides, extending through and transversely 
subdividing the fin-muscles and joining and blending with the 
fin-ray. The same disposition is observable in other fishes and 
is illustrated by Ceratodus (see p. 281). In the case of the 
pectoral and ventral fins the fascia is coittinued beneath the 
skin upon the muscular covering of the fins. 

The INTERNAL or ' TRANSVERSALis' FASCIA, lining the abdo- 
minal space and closely applied to the ribs, is distinct and 
strong. It is particularly strong in the region corresponding 
with that of the depressorea costarum of other animals. A thin 
layer of it bends inwards upon the tBsophagua. behind the peri- 
cardium, forming a rudimentary post-cardiac diaphragm. The 
fascia itself, or the greater part of the thickness of it, passes 
upon and is closely connected with the pericardium, and is, with 
the pericardium, attached strongly to the hinder and inner 
surface of the coracoid arch inferiorly. Superiorly it is thinner, 
and is accompanied upon the front of the pericardium by a 
distinct detachment of fibres from the internal surface of the 
ventral muscle, which thus forms a precardiac or post-branchial 
musculo-fascial diaphragm. Internally, the transversalis fascia 
is, in the greater part of its extent, in contact with the peri- 
toneum. Externally, it lines the ventral muscle and is con- 
nected with the external fascia by the intermuscular septa 
passing obliquely between them. Inferiorly, it passes beneath 
the edge of the ventral muscle and blends, as just stated, with 
the external fascia. Behind the pelvis it forms a sheath upon 
the bowel passing to the anus. (Fig. 23, 0,) 

The Ventral Muscle 

presents nothing remarkable in the tail. 

In the abdominal region, in about the middle third of the 
space between the lateral septum and the ventral mesial line 
(Fig. 23), the fibres of the outer stratum present a slight but 
distinct obliquity, from before backwards and cUyumyrAvdB (to- 

18—2 



256 PROFESSOR HUMPHRY. 

wards the mesial line), as they pass between the several s^ta. 
This disposition to the formation of an ohltquus extemus muscle 
is most marked in the middle of the space indicated. It dimi- 
nishes above and below; and in the third of the muscle next the 
lateral septum, as well as in the third next the mesial line, the 
fibres dixe* recti! The removal of the superficial — external oblique ^ 

— stratum shews an obliquity, to a like degree, but in a different 
direction, in the subjacent stratum. (Fig. 23 A) The fibres 
there pass from before backwards and t^pwards (towards the late- 
ral septum) and form an ohliquus intemus. In this instance also 
the oblique direction disappears in the upper and lower thirds 
of the muscle. The fibres there run antero-posteriorly, and the 
tendency to division into planes is lost. This gentle effort, 
as it were, at the formation of oblique muscles is an interesting 
corroboration of the remarks made (p. 11) with reference to 
the cleavage of these muscles in Cryptobranch, exhibiting as 
it does a still lower grade of stratification than we found in 
that animal (See also Ceratodus, p. 280.) There is no ap- 
pearance of a third or transversalis stratum ; and there are no 
muscular fibres in a deeper plane than the ribs. This is also 
the case in most Fishes, though in some (Bream and Dace) there 
is a thin muscular layer beneath the level of the ribs; and the 
nerves lie upon its outer surface. 

In describing the external fascia, I have mentioned that a 
superficial layer of it is continued upon the exterior of the 
mesial fin and that another layer passes beneath it; so that the 
rays and muscles of the fin are enclosed between the two layers 
and are separated from the lateral muscle by the deeper of 
them: also that the intermuscular septa are continued into the 
fin and blend with the rays, so causing a transverse segmenta- 
tion of the fin. The fin-muscles are contained in the fascial 
partitions thus made, and consist of a layer of fibres paraUel 
with the fin-rays, resting with one end upon the lateral muscle, 
or rather upon the fascia which separates them from it, and 
with the other end expanded upon the fin. The greater number 
of the fibres arise from and lie along the lines of the septa and 
the rays, as shewn in Fig. 23, where the alternating thicker and ^ 

thinner parts of the muscle thus produced are well displayed, and 
the thicker parts are seen to be in or near the lines of the 



MUSCLES OF LEPmOBIREN. 257 

septa of the lateral muscle and of the fin-rays. Now, the septa 
are disposed with regard to the fibres in the same way as they 
are with regard to the lateral muscle; and they connect the 
divisions of the fin-muscle with one another, and with the lateral 
muscle, much in the same way that they connect the several 
divisions of the lateral muscle itself. The chief difference 
consists in the direction of the fibres of the fin-muscle, which 
is at right angles, or nearly so, to that of the fibres of the 
lateral muscle. This causes the one series to be segmented from 
the other; and the action of the two being distinct leads to 
their being further separated by the fascial layer passing be- 
tween them. At the forepart of the dorsal fin the fibres of the 
fin-muscle extend a short distance, on either side, upon the 
surface of the lateral muscle between the two layers of fascia. 
They do this more widely in some Fishes, becoming thus sub- 
cutaneously expanded, upon the dorsal muscle; and they are 
still more segmented from it\ 

The relations of the ventral muscle to the ventral-fin are 
simple and interesting. The chief part of the muscular fibres of 
the fin are derived from the slanting poetero-lateral edge of the 
pelvis', above and below the acetabulum, and indeed all round 
the joint. These (Fig. 23 E) descend upon the whole circumfe* 
rence of the fin, ensheathing it and blending with the fascial 
tissue beneath the skin and with the fibrous perichondrial tissue 
which invests and unites the series of cartilaginous pieces that 
form the cylindrical tapering skeleton of the fin. To these 
are superadded fibres from the adjacent part of the ventral 

^ See, farther, the description of the relation of the fln-mnsoles to the 
lateral mnscle in Ceratodas (p. 280). 

' The pelvis is of somewhat diiunond shape. The posterior angle is pro- 
longed slightly into an obtusely pointed process. The anterior angle runs for- 
wards as a smooth styliform process, an inch and a half long, and occupies a 
channel in the fascia formed in the same manner as the channel for the mesial 
fin above described, a circumstance which is not a little confirmatory of the 
view I have expressed (Jvam. ofAnat. v. 69) respecting the serial homologous re- 
lations of the mesial and lateral fins. The lateral angles of the diamond run 
out into horn-like processes, and are connected with or continued into the 
transverse septa of the lateral muscles on the two sides. Near the hinder part 
of the cartilage, on either side, is the articular facet for the first cartilage of the 
fin. See Fig. 28. 

In Owen's Memoir, Linn. Tram, xvm. Tab. 23, as well as in the Anat. Vert, 
I. Fig. 47, the pelvis is represented and described with the styliform process 
directed backwards. 



258 PROFESSOR HUMPHRY. 

muscle, which coa verge from before and from behind and blend 
with them. There is no line of demarcation between the fibres 
derived from the pelvis and those derived from the ventral 
muscle. Some of the latter proceed from the next adjacent 
septa before and behind; and some are derived from more 
distant septa. As represented in Fig. 23, the lowest and hind- 
most of the ventral fibres passing upon the fin are segmented 
from the rest of the ventral muscle and are uninterrupted by 
septa for a greater distance than those situated higher up. 
Between the converging bundles from before and behind is seen 
an interval (F) in which the nerves run, from beneath the edge 
of the ventral muscle, over the outer edge of the pelvis and the 
acetabular joint, upon the fin. 

As it approaches the forepart of the animal, the ventral 
muscle (and this will be found to be the case also with the 
dorsal muscle) gives off a superficial stratum which is thin and 
partly membranous (Figs. 24 and 26). Its disposition is as 
follows: it separates from the deeper stratum of the ventral 
muscle a little behind the level of the pectoral fin, at one of 
the septa, and is thenceforward connected with the deep stratum 
only by loose areolar tissue; so that it is easily dissected away. 
This is especially the case near the middle line. It retains the 
transverse septa and, advancing forwards, expands as a con- 
tinuous sheet upon the shoulder-girdle and the hinder aspect of 
the base of the fin. It is also continued forwards upon the 
lower jaw, the gill cover and the opercular bones; and it 
spreads in this direction upon the surface of the dorsal muscle, 
becoming blended with the fascia of that muscle and so reaching 
the dorsal median line. It may be called the SUPERFICIAL 
BRACHio-CEPHALic STRATUM of the ventral muscle, to distin- 
guish it from the deeper stratum of the same muscle which also 
passes to the limb-girdle, limb and head, and which may be 
called the deep brachio-cephalic stratum. 

The upper portion {L. d.) of this superficial brachio-cephalic 
stratum — the portion, that is, nearest to the lateral septum, — a 
little in front of the line of separation from the deep stratum, 
is inserted into the upper, or scapular, part of the shoulder- 
girdle, which forms a conical projection above the fin, and into 



MUSCLES OF LEPIDOSIREN. 259 

the upper part of the first cartilage of the fin, which may be 
regarded as representing the upper or humeral segment of the 
forelimb of higher animals. Some of the fibres also blend with 
the other muscles of the fin. This portion of the stratum 
represents, therefore, the latissimua dorsi. 

The next portion (jP) of the stratum, a little more ventrally 
situated, is attached to the under-surface of the fin and its first 
cartilage, and to the anterior edge of the shoulder-girdle (the 
coracoid) beneath the fin. The coracoid is here covered by an 
extension of the mucous membrane of the branchial chamber, 
and contributes to form the wall of the branchial passage, and 
of the slit-like branchial opening; and the mucous membrane 
extends a short distance upon the surface of the muscle which 
accordingly contributes, with the coracoid, to form the hinder 
wall of the passage leading to the branchial chamber. This 
portion of the stratum corresponds with the pectoralia major. 
It is really continuous with the latissimus dorsi: but the fibres 
establishing the continuity between the two, and attached to 
the girdle and the fin in the immediate neighbourhood of the 
glenoid cavity, form a very thin sheet; and an axillary interval 
is easily made between the pectoral and the latissimus dorsi 
portions of the stratum. 

Still more ventrally or mesially, the fibres of the superficial 
brachio-cephalic stratum pass onwards and form a superficial 
cervical muscle (cervicalis swperficiaUa). The lateral fibres run 
beneath and in front of the branchial opening ; and, covered by 
the skin externally and lined by the mucous membrane inter- 
nally, they form the margin of the gill cover and the anterior 
wall of the passage leading to the branchial chamber. They 
(Fig. 24, Sp. 6r.) moreover spread out over the branchial cham- 
ber and, in front, come into contact with and are attached to 
the opercular bones ^ and the side of the skull; while, above, 

^ There are distioctly two opercular bones as reproBented by Petem. One, 
tbe inferior or titbopereulum, sitoated behind the an^e of the mandible, is tri- 
angular, with its apex directed forwards and continued into a narrow tendon 
wluch is attached to the inner surface of the lower part of the dentary bone, 
about the middle ; and it blends there with the fibres of the mylo-hyoid muscle. 
The other, the superior bone or operculum, is smaUer, more elongated, and 
extends up between the preceding and the hinder surface of the suspensorium. 

Owen (p. 336) mentions only the superior. Huxley, Anatomy of Vertebrate 
AmmalSy p. 170, mentions the superior as operculum and the inferior as a 
' branofaiostegal ray* carried upon the hyoidean arch. It is however oonneoted 



260 PROFESSOR HUMPHRY. 

they arch over the branchial opening and radiate into the fascial 
tissue covering the dorsal muscle (Fig. 24, Tr.). Here the 
hindmost fibres, bending backwards, are inserted into the 
upper and anterior surface of the scapula above the fin ; and 
some pass on into the fin. These returning fibres therefore 
meet the fibres of the latissimus dorsi advancing to the same 
part. They — so much of this part of the stratum as lies above 
the branchiae— appear to represent the trapezius^*, while the 
fibres lying below and upon the branchiae, and advancing for- 
wards to the skull, may be regarded as representing the stemo- 
deido-masfxnd*. Dissecting away the fibres which radiate 
upon the dorsal muscle, I find some beneath them, running 
below the level of the lateral septum and inserted into the 
supra-scapular bone, which is here deeply seated, also into the 
ligament which connects that bone with the scapula^ 

The dipping down of tfae deeper fibres of this layer, which I sap- 
pose to represent the trctpezius and atemo-mattoid among other 
muscles, towards the ceryical transverse processes accords with the 
disposition of the cervico-humeral muscle ia many mammals, and 
affords an explanation of that which has seemed to be an aberrant 
feature of this muscle, viz., that in its upper part it is in close rela- 
tion with the levator acapulce and might be thought to belong to it; 
whereas, below, it is in close relation with the trapezius and claims 
even more distinctly to appertain to it. It is farther interesting to 
remark that in the Crocodile the atemo-mcutoid runs to the cervical 
transverse processes; and a deep band of the trapezius does the 
same\ 

witLthe lower jaw and the operonlnm rather than with the hyoid; and its oper- 
cular relations are confirmed by the disposition of the corresponding bone in 
Ceratodus. This is described as suboperoalum by Gunther in his excellent 
paper on the Ceratodas, PhiL Trans. 1871, p. 525. 

^ It is named levator soapula by Owen, bnt is too superficially placed for that 
muscle. 

* These, at least, are the mnscles which they chiefly represent. There are, 
however, other muscles in different animals which must be derived l^ segmen- 
tation from the same source, such as the constrictor faucium, depressor inandi' 
bula, levator arcuum^ subcutaneus coUi, from the superficial surface, and, more 
deeply, the rhdmboids, masto-scapulnrt eervico-humeralf &o. 

* This bone is quite distinct, as described by Peters. For further account 
of fibres, similar to those above mentioned, see description of Dog-fish (p. 274). 

^ In the Dog-fish (p. 274) all the fibres pass beneath the lateral septum, and 
none expand upon the dorsal muscle. 

In that animal the several gilUopenings cause a greater separation between 
the upper and lower parts of the muscle than is caused by the single transverse 
elit'like opening in Lepidosiren, and I have accordingly named the part above 
the gill-openings cerviealis superficialis superior, and the part below them 
cervUaUs superficiaUs inferior^ 



K 



MUSCLES OF LEPIDOSIREN. 261 

The most ventral or mesial portion of the superficial brachio- 
cephalic stratum, separating a little from the portion last 
described, passes beneath the floor of the mouth and is inserted 
along the inner and lower edge of the circle of the lower jaw, 
also strongly to the comu of the hyoid^ near the augle of the 
jaw, and less strongly along the upper edge of the cornu, and 
to the suboperculum. Here it is blended with the subopercular 
fibres of the last described (stemo^leido^mastoid) portion. This 
portion of the stratum (Fig. 26, M. h. and 8. c.) represents the 
mylo-hyoid with part of the svbctUaneua colli or platysma. The 
transverse septa cease on a level with the angles of the jaw ; 
and the fibres in front of that take a more transverse direction, 
are less closely connected with the skin, and are less stained 
with pigment than in the hinder regions of the stratum. The 
stratum is here, for a short distance, in two layers (Fig. 26, S. c. 
and G. A.). In the deeper of these, which is a more direct con- 
tinuation of the superficial stratum of the ventral muscle, the 
fibres have an antero-posterior direction. It is inserted near the 
symphysis of the jaw only ; and just behind the symphysis it is 
in immediate contact with the mucous membrane of the 
mouth, which is reflected upon it from the rudimentary hyoid 
and tongue and then passes from it to the symphysis. In the 
superficial layer (8, c.) the muscular fibres are more transverse, 
and are inserted into the ramus of the jaw and the comu of 
the hyoid as well as into the symphysis. It is this superficial 
layer which more distinctly represents the myh-hyoid and the 
svbcutaneus coUi, while the deeper layer {6. h) may represent, 
in part, the genio-hyoid^ ; but the two layers are blended in 
front, near the symphysis, as well as behind at their junction with 
the septum of the ventral muscle from which they both spriug. 

It should be added that some of the fibres of the superficial 
layer, lying between those which I call avhcntaneus colli and 
those which I call mylo-hyoid, radiate from, as well as are 
inserted into, the angle of the jaw and the adjacent comu of the 
hyoid ; and the fibres passing from the latter point, beneath the 
body of the jaw, to the symphysis form a hyo-mental (p. 325) and 
appear to represent the anterior belly of the digastric (Fig. 26, D). 

1 The oerato-hyoid. There is no hasi-hyoid. 

* They are so named by Owen in the desoription of Figs. 4, 5, and 6. 



262 PBOFESSOR HUMPHBT. 

The DEEP BRACHIO-CEPHALIC STBATUH of the VENTBAL 

MUSCLE (Fig. 26, D. v)i the pai*t that is beneath the whole of 
the superficial stratum just described, is far thicker than it and 
is marked by septa in its whole length. In its course it en- 
counters three bones in succession ; each of which is in the 
line of one of its septa and is involved in it. 

The first of these bones (Fig. 26, A) is a stout cylindrical 
elongated bone, articulated by a broad trochlear surface with 
the cartilage of the base of the skull just behind the opening 
for the vagus nerve, and running down into the substance of 
the lateral muscle. This bone in form, position and relation 
to a septum, resembles a rib; and it probably is to be regarded 
as a rib, although it is much larger than any of the succeeding 
ribs. This view is confirmed by the fact that a short thick 
muscle here separates itself from the rest of the deep stratum 
of the ventral muscle and passes from the bone in question to 
the under surface of the scapular part, of the shoulder-girdle, 
representing pretty clearly the serraiua^. The nerve to the 
fin takes its course beneath the edge of this muscle, between 
it and the muscles attached to the coracoid, holding therefore 
the same relation to it as the brachial nerves in other animals 
do to the serratua. 

This deep stratum of the ventral muscle next comes into 
contact with the coracoid which is connected with one of its 
septa, much in the same manner as the ribs are connected with 
the septa of the ventral muscle, that is to say, the coracoid is an 
ossification in the deepest part of the septum. Many of the 
muscular fibres are inserted into it, representing the costo- and 
stemo-coi^acotds (the ribs and costal cartilages being represented 
by the septa behind the coracoid). The greater portion of the 
fibres, however, run beneath, superficial to, the coracoid on to 
the hyoid, constituting a deep cervical {cervicalis profundus/) 
muscle (C p.). Some of these are inserted, tendinous, into the 
hyoid near the middle line. Some terminate in a tendinous 

^ I am glad to find that the yiew I have formed of this peculiar bone from 
its moscnlar relations in Lepidosiren is confirmed by Gunther, in his description 
of the corresponding bone in Ceratodus, Phil. Trans. 1871, p. 529. Various 
views haye been entertained of it, the last being that by Mr Parker (Shoulder' 
girdle and Sternum, p. 21, and PI. n. Fig. 1), who regards it as a 'pharyngo- 
branchial' bone, and a small cartilage attached to its middle as a * small onos- 
sified second phaiyngo-branohial.* 



MUSCLES OF LEPIDOSISEN. 263 

band which is continued round the anterior part of the hyoid, 
near the middle line, and is inserted into the dense tissue 
there, in the plctoe of a glosso-hyal, which causes a median 
elevation in the floor of the mouth, behind the symphysis of 
the jaw, and constitutes a rudimentary tongue. The more 
lateral fibres, curling beneath the coracoid, are folded over the 
median part of the muscle between the coracoid and the hyoid, 
giving a thick rounded edge to the muscle in this situation, and 
are inserted into the upper surface of the middle of the hyoid. 
The result of this disposition is a thick strong coraco- or ventro- 
hyoid, a representative of the stemo^hyoid, which serves power- 
fully to draw the hyoid backward, deepening the cavity of the 
mouth and raising the gill-cover. To some extent, therefore, 
it acts antagonistically to the superficial stratum, part of the 
office of which is to compress the opercular apparatus. It is 
disencumbered from other muscles by complete segmentation, 
and by an investment of loose areolar tissue, so as to be enabled 
to act freely and fully upon the hyoid. 

The pharyngeal muscles of higher animals are represented, 
or partly represented, by a delicate layer of fibres — a Ayo- 
pharyngeus — passing backwards firom the hyoid, on either side of 
the mesial line, and radiating upon the mucous membrane of the 
pharynx. They meet and blend with a similar layer of fibres pass- 
ing forwards from the coracoid and, in like manner, radiating upon 
the pharynx and the under-surface of the branchial chamber. 
These sub-mucous layers, specially segmented from the deep 
surface of the ventral muscle, resemble and are derived in the 
same manner as the subcutaneous layers which are specially 
segmented from its superficial surface, and which constitute 
the pUUymna and the panniculus camosus of Mammals. 

A short strong fan-shaped muscle — stylo-hyoid — radiates 
from inner surface of the hyoid \ near its upper end, to the 
under surface of the back part of the suspensorium, behind the 
auditory capsule, where it has a broad insertion. 

^ The hyoid (cerato-hyoid) reste npon the.anditoiy oartilage, and is con- 
nected with the base of the sknll by a tongh ligament, and with the inner side 
of the angle of the jaw by a strong stylo-, or better, hyo-maziUaxy ligament. 



264 PROFESSOfi HtJMPHRT. 

Dorsal Muscle. 

The superficial stratum of the dorsal muscle, separating in 
the same manner and nearly at the same part of the animal 
as that of the ventral muscle, soon becomes in great part 
membranous, passes over the side of the head, and is attached 
to the end of the suspensorium just above the glenoid cavity, to 
the lower jaw a little in front of its angle, to the tooth-like hinder 
projection of the maxillary bone and to the edges of the pre- 
maxillary and frontal bones ^. It represents the temporal fascia; 
and muscular fibres are attached to or detached from both its 
surfaces. Those which are connected with its superficial surface 
are continued into, or are more or less connected with, the 
superficial stratum of the ventral muscle. It is thus connected 
with the muscles already described as representatives of the 
trapeziua and stemo-masixiid ; also with the two following 
muscles situated in front of those last-mentioned and, like 
them, appertaining rather to the superficial ventral stratum; 
though their description is more conveniently given here. 

These two muscles are superficial and thin, and lie upon 
the extension of the dorsal fascia to the suspensorium and 
mandible. They are separated from one another only by a 
fascial line from which the fibres of both arise; so that they 
might be regarded as one. The hinder of the two, which I will 
call levator operculi (Fig. 24, L. o.), radiates, from the angle of 
the mandible and the fascial line just alluded to, backwards 
upon the superior opercular bone and the interval between the 
two opercular bones where it blends with the pre-branchial 
(stemo-cleido-mastoid) part (^. 6r.) of the ventral muscle. The 
anterior of the two muscles (Fig. 24, D, m.) is also attached 
to the angle of the mandible, to the fascial line and to the 
suspensorium. Its fibres are directed more forwards upon the 
dorsal fascia. Its position is not unlike that of the masseter ; 
but the masseter is represented by another muscle. On the 
whole, it rather corresponds with the depressor mandibulce, or 
with that portion of it which, in some animals, arises from the 

1 MtUler (8. 245 and Tab. ti.) describeB in BdeUostoma a similar extension 
of the dorsal musole forwards over the side of the head to the tentacles. It 
was connected by a ligament with the hyoid and the npper part of the sknlL 



MUSCLES OF LEPIDOSIREN. 265 

Buspensorium; while the portion which, in Cryptobranch and 
many others, arises from the dorsal fascia and the cervical 
spines is probably represented, in Lepidosiren, by some of the 
fibres of the levator operculi and of the supra-branchial (tra« 
pezius) muscles. 

One or two points deserve attention here. Firaty the mode in 
which the superficial and the deep strata of the dorsal and of the ven- 
tral muscle are blended, rendering it difficult or impossible to say 
where the one begins and the other ends, or to refer precisely the 
respective muscles to the one or the other. This blending seems, on 
the whole, to result from an extension of the ventral upon the dorsal 
stratum; and we find the same thing often occurring, behind the fore 
limb, iu the spreading of the latissimus dorsi towards, or to, the verte- 
bral opines. Secondly y the superficial and the deep strata, ventral and 
dorsal, are blended respectively in the ventral and dorsal muscles be- 
hind; both sti-ata may contribute to the formation of the same muscle, 
as we shall presently find in the case of the temporal muscle. Thirdly, 
the superficial pre- and supra-branchial stratum which we have been 
considering as a derivative chiefly from the ventral muscle, presents 
itself in Lepidosiren in a simple, but from its connections very sug- 
gestive form, warranting, I think, the view I have taken that it is the 
representative, not only of the trapezius and atemo-mastoid, but of 
the numerous other muscles segmented in different ways in different 
animals, and passing upon the shoulder, the neck, the branchial 
apparatus and the jaw, so as to meet their varied requirements. 
F&wrMy^ it is interesting to note, in connection with the relation of 
the superficial strata of the lateral muscle in other parts and other 
animals to the so-called membrane bones, that the snper6cial stratum 
of the dorsal muscle 5s in Lepidosiren blended with the periosteum 
of the membrane bones of the upper and anterior regions of the skull. 
Further, some of these bones partially enclose the eye and form the 
orbit^ fi'om which, or its periosteum, the muscles of the eye arise. 
The ocular muscles in Lepidosiren surround the optic nerve. They 
may be clearly distinguished as four recti muscles of which the 
external is the largest; they are distinctly tituseable to the peri- 
osteum in which, as just said, the dorsal stratum ends; and they are 
thus brought into continuity with the superficial stratum of the 
dorsal muscle and shewn to be derivatives from, or prolongations 
o^ this part of the great lateral muscle. 

The reflection of the superficial stratum brings into view 
the DEEP DORSAL STRATUM which is thick and composed of 
fibres passing forwards to the skulL A partially segmented 
conical portion of it, commencing about the third vertebra, 
and expanding as it advances forwai'ds, forms the iem^porol 
muscle (Fig. 25, T). It passes over the suspensoriam, filling 



266 PROFESSOR HUMPHRY. 

up the lateral concavity of the skull. It is covered by the 
superficial stratum (the temporal fascia), as well as by the 
overhanging cranial bones into which that stratum is inserted. 
It derives a large accession of fibres from these structures 
which form its covering, as well as from the parts of the skull 
upon which it lies. It constitutes the hinder wall of the 
orbital space; and its fibres converge to be inserted into the 
coronoid process of the mandible, which is well developed, rising 
up behind the dentary plates that cover the forepart of the edge 
of the jaw. 

The large size and extensive attachments of this muscle correspond 
with the thickness of the mandible, with the secure mode in which it 
is articulated with the skull and with the strong dentary plates which 
it carriea Its relation to the dorsal muscle, foreshadowed in Crypto- 
branch (PI. rv. Fig. 20), and clearly made out in Lepidosiren, is of 
great interest as establishing the direct continuity of the muscles of 
the jaw with the lateral muscle. 

The under surface of the temporal muscle, at and in front of 
the suspensorium, is confluent with the masseter. The division 
between the two is, however, indicated by the passage of the 
branches of the third division of the 5th nerve ; and the latter 
muscle, though really a continuation of the temporal, may be 
described as arising from the forepart of the suspensorium and 
the inner surface of the tempoi-al fascia ; and it is inserted into 
the outer side of the coronoid process, beneath the temporal 
muscle, and into the outer side of the jaw near the coronoid. 

Muscular fibres, arising from the forepart of the coronoid 
process, and partly continuous with those of the temporal muscle, 
radiate forwards upon the mucous membrane of the mouth, 
beneath the retractor anguli oris, and constitute a byccinator. 

A stout retractor anguli oris (Figs. 24j and 25, R. a, o.) arises 
from the extremity of the suspensorium and the adjacent inner 
surface of the temporal fascia, also by a strong tendon from the 
outer side of the mandible in front of the angle. It is lost in 
the tough tissue about the angle of the mouth. It appears to 
represent in part the depressor anguli oris and perhaps the 
Zygomatid. 



NERVES OF LEPIBOSIBEN. 267 



i 



CEREBRAL NERVES. (Fig. 27, alao Pig. 25.) 

I subjoin a short account of the cerebral nerves of Lepidosireny 
disseotecl by Mr Anningson. 

The Optic nerve emerges from a small foramen in the forepart of 
the side of the skull beneath the frontal bone. It is accompanied and 
surrounded by the four recti muscles. Special nerves to these muscles 
(the third, fourth, and sixth) were not found. 

The FIFTH nerve passes, in three divisions, through three foramina 
situated in the cartilage forming the anterior waU of the skull beneath 
the edge of the squamous bone. 

The uppermost or ophthalmic division (Op?Uh,) passes, deep, along 
the inner wall of the orbit, crosses over the optic nerve, gives off ciliary 
and oculo-motor nerves, and divides into three branches at least. One 
(/ronto-'iuisaly /. n,) is joined by a branch from the supra-maxillary, and 
is distributed upon the inner side of the nasal process of the skull, 
upon the surface of that process, and upon the adjacent part of the 
head. A second branch (nascd^ n.) traverses the inner wall of the 
orbit to the nasal cavity. A third branch reaches the floor of the 
orbit and unites with a branch of the supra-maxillary to form the 
in/rcHyrbUal nerve (i. o.) which is distributed in the upper lip. 

The SUPRA-MAXILLART division {S. m.) gives off three chief branchesy 
a 'fronto-nasal,' an * infra-orbital ' and a ' muscular/ The /ronUhnasal 
{/' **'•) piercea the upper part of the temporal muscle (Fig. 25/. n.\ 
and runs upon its suHace, beneath the temporal fascia and beneath 
the edge of the frt>nto-nasal process. There it joins the fronto-nasal 
branch of the ophthalmic divinion. The second or in/ra-orhital branch 
(t. o\) runs beneath the maxilla just in front of the orbit and joins 
the infra-orbital branch of the ophthalmic. The third, or musctdar^ 
branch (t.) passes into and is lost in the temporal and masseter 
muscles, and appears to be the only nerve to those muscles; at least 
no branches could be traced into them from the next division. 

The infra-maxillary division escapes from the skull in two 
branches separated by membrane but traversing one foramen. One 
of these gives off a nasal nerve (n.) which pierces the temporal 
muscle and runs beneath the temporal fascia (Fig. 25 n .), passes to the 
nose, partly joins the nasal branch of the ophthalmic and is partly 
distributed to the cheek. This branch gives off a buccal nerve (Figs. 
25 and 27 b,) which is also distributed to the cheek. The other, or 
mandibular, branch (m.) of this division is the larger of the two. It 
inclines downwaixls and forwards, passes over the upper edge of the 
mandible, between its coronoid and articular parts, runs beneath the 
strong external ligament of the joint and between the temporal and 
masseter muscles. Having gained the outer sur&oe of the jaw it 
divides into three nerves. Two of these (Figs. 25 and 27 1,) are labial 

^ This mosoolar branch, as wall as the infra-orbital, is in Cryptobranoh de- 
rived from the infra-mazUlaiy (p. 46). 



2G8 PROFESSOR HUMPHRY. 

branches supplying the integuments on the exterior of the jaw and lip ; 
and the third penetrates the jaw between the dentary and the articu- 
lar parts. 

The SEVEirrH nerye traverses a hole in the cartilage on the under 
sur&ce of the skull, near the forepart of the suspensorium, in front 
of the origin of the stylo-hyoid muscle. It runs outwards between 
that muscle and the skull, gives off branches to the stylo-hyoid and 
to the muscles upon the opercular bones. One large branch (7') 
passes forwards between the suspensorial cartilage and the squamous 
bone, meets the labial branch of the infra-mazillary, accompanies it 
over the edge and upon the outer surface of the mandible, and blends 
with one of the branches of it going to the lower lip. Another 
large branch (7") passes beneath the suspensorium, supplies the de- 
pressor mandibulse, perforates the strong stylo- mazOlary ligament, 
and runs external to the ligament connecting the inferior opercular 
bone with the mandible. It continues its course under the shelter of 
the dentary bone, close to the mucous membrane, between it and the 
mylo-hyoid. Finally (7'"), it pierces the mylo hyoid, curls over the 
edge of the jaw, and blends with the lower of the two infra-maxillary 
branches to the lip. 

Peters represents this as the third division of the 5th. Its 
foramen is in front of the usual position. Still it is quite beneath 
the skull; its branches are distributed much in the same manner as 
those of the seventh nerve in Cryptobranch (p. 46), and the three 
divisions of the 5th are quite distinct in Lepidosiren. 

The blending of the branches of the several divisions of the fiflbh 
with one another, and of the branches of the infra-maxillary division 
with those of the seventh — ^their imperfect isolation — ^is of much interest^ 
in connection with the want of isolt^tion of the oculo-motor nerves in 
these animals. It is also interesting in relation to the union which 
in ourselves is established between the supra- and infra-maxillaries and 
the seventh through the medium of the corda typani and other petro- 
sal nerves. The object, whatever it may be, which is attained by the 
blending of the branches in these animals is in ourselves effected, 
apparently in part, through the medium of the sphenopalatine and 
otic ganglions. 

It is remarkable that the supra-maxillary supplies the chief if not 
all the branches to the temporal and masseter musclea The upper 
portion of the infra-maxillary, though passing through the same fora- 
men with the rest of that division, corresponds with branches of 
the supra-maxillary division in other animals. 

The EIGHTH nerve or Yagus emerges from a hole in the aide of 
the skull behind the suspensorium, the auditory cartilage being 
between it and the seventh nerve. It immediately breaks into 
branches. Of these some pass to and supply the branchiae and some 
pass to the pharynx. One of the branchial nerves (that to the 5th 
branchia) runs along the branchial arch to near the middle^ passes 
between the pericardium and the coracoid, penetrates the ventral 
muscle and runs backwards, beneath the skin, near the ventral 
mesial line. A eomco-hyoid nerve runs downwards, behind the 



NEBVES OF LEPIDOSIREN. 269 

bnmchuey nearly parallel to the preceding, and enters and is distri- 
buted in the deep stratum of the ventral muscle between the coracoid 
and the hyoid. The Fneumo-gastric passes backwards to the lungs 
and along the ossophagus. The Lateral nerve is the largest diviHion 
of the vagus. It sends a branch upwards, behind the suspensorium 
to the deep part of the dorsal muscle. It then runs directly back- 
wards upon the dorsal surface of the peculiar bone (the large first rib) 
and the other ribs, and the lateral septum, continuing its course 
immediately beneath, or in the deepest part of the substance of, the 
dorsal muscle. 



DESCRIPTION OF FIGURES. PI. XII. 

Fig. 23. The part of the animal in the neighbourhood of the 
ventrtd fin& The skin and the fascia have been removed to shew the 
dorsal and ventral museles with the lateral septum between them. 
The fibres of the ventral muscle, midway between the lateral septum 
and the mesial line, are oblique from before downwards and back* 
wards. Those of the subjacent stratum (exposed at A by dividing 
the superficial stratum along the interval between two of the 
transverse septa and reflecting it forwards and backwards) are, in the 
same region, oblique from before backwards and upwards. £, the 
. edge of the fascia seen to split at the base of the dorsal fin. C, the 

( muscles of the dorsal fin thickest in the neighbourhood of the trans- 

verse septa, and these overlying the raybones. D, the pelvic carti- 
lage with its anterior styliform process lying in a facial sheath, and 
its ascending lateral horn. ^, ventral muscle continued in front 
and behind upon the fin. F, nerves converging upon the dorsal 
aspect of the fin. &, lower part of the intestine ensheathed by the 
fascia transversalis and terminating in the cloaca which is on the left of 
the mesial line. (It is usually on one side or the other in this animal, 
as observed by Peters Mailer's Archiv, 1845, p. 9. It was on the 
left side in Owen's specimen.) The muscles on the anal fin resemble 
those on the dorsal fin. 

Fig. 24. Superficial layer of muscles on left side of fore part of 
body, fin and head. J)^ superficial stratum of dorsal muscle con- 
tinued into temporal fascia which is continuous with frontal bone (F). 
This &scia has been cut away in front where it was attached to (A^) 
the nasal bone, (Fm.) the premaxillary bone, (N, c.) the nasal carti- 
lages, {M) the maxillary bone, (De,) the dentary part of the lower jaw, 
and (An,) the angular part of the lower jaw. T^ temporal muscle. 
F, a, 0., retractor Anguli oris. L. d:, latissimus dorsi. F, pectoral. 
^. 6r., supra-branchial, or cervicalis superficialis, portion of ventml 
muscle. Behind the last is the slit-like branchial opening, which is 
bounded behind by the coracoid with the fibres of the ventral muscle 
passing to its edge and the coraco-brachial fibres ((7. br.) running fiom 
its edge upon the fin. Between the latissimus dorsi and the pectoral 

VOL, VI. 19 



270 PEOFESSOR HUMPHRY. NERVES OF LEPIDOSIREN. 

is the axilla. 5V., trapezius. 0* and 0', inferior and superior opercu- 
lar bones. L, o,, levator operculi. D, m., depressor mandibulae. 

Fig. 25. Similar view to the preceding in which the superficial 
dorsal layer (D. D.) and the temporal fascia have been divided longi- 
tudinally and reflected upwards and downwards exposing {T) the 
temporal muscle. My the masseter with the branches of the third 
division of the fifth nerve emerging from between it and the temporal 
muscle. R, a. o,, retractor angiili oris, arising from the suspensorium. 
Fj frontal bone. Pm,, premaxillary bone. F. n\, fronto-nasal branch 
of the supra-maxillary division of the fifth nerve; t. o'., infi'Srorbital 
branch; n\ nasal branch of infra-maxillary; ly labial, and m, man- 
dibular branches of the infra-maxillary. 

Fig. 26. Deep dissection of throat showing disposition of ventral 
muscle. The superficial stratum (S. c.) of lighter colour has been 
partly divided on the left side and reflected towards thejaw, where, on 
its under surface M. h., the mylo-hyoid, is seen attached to the jaw and 
the hyoid, and also forming (D) the anterior pai*t of the digastric. 
G. &., coraco-brachial. N, nerve passing to the ventral aspect of the 
fin. Ff pectoral. M. m,^ mucous membrane reflected from hyoid 
and rudimentary tongue upon superficial stratum of ventral muscle. 
S, c. and G. A., the two layers of the superficial stratum of ventral 
muscle. G, p., cervicalis profundus, or deep stratum of ventral muscle 
between {G) the coracoid and the hyoid. Ry thick rib-like bone 
exposed by dividing part of deep stratum of ventral muscle. 

Fig. 27. Kight side of skull of Lepidosiren, from which the 
muscles have been cleared away to show the nerves. The optic is 
not shewn. Opluh., the ophthalmic of the fifth;/, n., its fronto-nasal 
branch; n^ its nasal branch; and f. o, its infra-orbital branch. S, m., 
the supi*a-maxillary division; t, its temporal or muscular branch; 
/in., its fronto-nasal branch; i, o\, its infra-orbital branch. /. m., 
the infra-maxillary division ; n , its nasal branch ; b, its buccal branch ; 
m, its large mandibular branch giving off (I) labial branches to outer 
side of mandible. 7\ the branch of the seventh accompanying the 
mandibular branch of the fifth; T\ the branch of the seventh to 
the depressor mandibulss (/>. nh), which is continued beneath the 
mylo-hyoid and curls beneath the jaw reappearing {7'") to supply the 
lip. Sq.y the squamous bone. S. o., the supra^^percnlar bone. /. o., 
the infra -opercular bon& G. A», the coraco- hyoid muscle. D, m., 
the depressor mandibulse. X, the external lateral ligament of the jaw. 



THE MUSCLES OF THE SMOOTH DOG-FISH (Mustelus 
levis). By Professok Humphey (PL XIIL, Figs. 28 
to 33). 

Figure 28 represents the dorsal aspect of the fore part of the 
animal, and shews the obliquity of the transverse septa in the 
upper half {M. D,) of the dorsal muscle, the mesio-dorsal 
muscle as it is sometimes called. In the lower or latero-dorsal 
half {L. D,) of the dorsal muscle, the septa form a series of 
gentle curves with anterior convexity; but in the upper half 
they are projected forwards into angles, which increase to ex- 
treme acuteness as the muscle approaches the head, more so 
than could be represented in the drawing. The upper lines of 
the angles are the more oblique and larger, running twice as 
far backwards as the lower lines or more. There are also a few 
return septa near the dorsal edge; and the angles formed by 
these, with the upper lines just mentioned, are prolonged back- 
wards into extremely fine points which are lost in the mesial 
septum. The pointed extremities of the several angles formed 
by the union of the septa are thicker than the rest of the septa 
and show a gradual transition into tendons, some of which are 
fully developed and run along as distinct delicate tendons for 
considerable distances without any accompanying muscular fi- 
bres; and they are inserted into the median dorsal septum or 
into the vertebral processes lying in or near the septum. These 
prolonged tendinous apices of the converged and united septa 
closely resemble the delicate tendons of the dorsal muscle so 
remarkably developed in Snakes, and are obviously the repre- 
sentatives of them, rendering it clear that the tendons and the 
septa are homologous structures. 

The muscular fibres situated between the very oblique septa 
do not retain their antero-posterior direction, but run with some 
obliquity in a penniform manner. 

Turning attention to the ventral muscle, a broad median 
portion — the medio-ventral portion (if. F.), of paler colour than 
the rest of the lateral muscle, is seen to overlap the more la- 
teral — the latero- ventral — ^portion, covering it as far as the in- 

19—2 



272 PROFESSOR HUMPHRY. 

verted angles formed by the ventral septa. It has a defined 
edge and is easily separated from the lateral portion, there being 
a distinct, though delicate, layer of areolar tissue between the 
two. This at least is true as regards the muscular fibres, which 
in both portions are throughout directed antero-posteriorly. 
The intermuscular septa are, however, continuous and hold the 
two portions in close relation with each other; and behind, near 
the ventral fin, the two are blended in the common ventral 
muscle. In the drawing the middle portion is shown partially 
reflected ; and the subjacent lateral portion is seen in shade. 

The median portion, which corresponds with the brcichio- 
cephalic portion of the lateral muscle, superficial and deep, of 
Lepidosiren (p. 258), increases in breadth and distinctness as it 
advances forwards, and is inserted into nearly the whole length 
of the shoulder-girdle and into the fin. A little behind the fin 
it gives off a superficial stratvum, which, however, is much less 
segmented from the deep stratum than is the corresponding 
structure in Lepidosiren. The upper portion of this stratum, 
representing the latisaimus dor si {L, d.), is inserted into the 
scapular part of the girdle, and expands upon the dorsal surface 
of the root of the fin reaching to its anterior edge. Upon the 
fin it lies upon, and to some extent blends with, the proper 
muscle of the fin. In the drawing it is represented as partly 
reflected from the fin-muscle. The lower portion of the stratum, 
representing the pectoralis (P), is inserted into the coracoid 
part of the girdle, and extends slightly upon the tmder or ven- 
tral surface of the fin, becoming, in some measure, blended with 
the proper muscle of the fin. An intermediate or axillary por- 
tion of the stratum forms a furrow, in which the edge of the fin 
is received. It is inserted into the girdle and into the fin, be- 
hind the joint of the fin with the girdle. 

I may here remark that the upper or scapular end of the 
girdle is prolonged into a pointed process, which runs over the 
lateral septum and projects in between the fibres of the dorsal 
muscle, where it terminates in a ligament which is lost in one 
of the transverse septa, not far from the lateral septum. I do 
not, however, find that any of the fibres of the dorsal muscle are 
inserted directly into the scapula. 

Before reaching the lateral septum and the dorsal muscle 



MUSCLES OF THE SMOOTH DOG-FISH. 273 

the scapula lies upon the upper part of the ventral muscle— the 
latero-veutral muscle — and its lower edge is connected with one 
of the transverse septa of this muscle. Moreover, some of the 
fibres of the muscle pass into its under surface, constituting a 
serrdtus. These are continuous with fibres of the same muscle 
farther forward, which pass, from a deep level of the lateral 
septum, backwards to the under surface oi the scapula, and are 
inserted there, constituting a levator scapuice. 

Beverting to the pectoral part of the mesio-lateral muscle, 
which I have described as being inserted into the coracoid, I 
should add that the more superficial fibres (Fig. 29) are inserted 
not into the coracoid but into the transverse septum lying be- 
tween the coracoid and the skin (the coracoid is developed in 
the deeper part of this septum, which may therefore be called 
the * coracoidal* septum). Through the medium of this septum 
it is continued onwards into a superficial cervical muscle, which 
may be called * cervicalis superficialis inferior* This last is a 
thin muscle without septa; and its fibres radiate forwards and 
outwards. Anteriorly, they pass superficially to the hyoid, ac- 
quiring reinforcements of fibres from the mesial line, and are 
attached to the lower margin of the mandible and also to the 
hyoid near the angle of the mandible. Posteriorly, they ex- 
pand, to some extent, upon the fin, as well as are continuous 
with the pectoral Laterally, they pass beneath the branchial 
chamber, are connected slightly with the branchial cartilages, 
and extend into the gill-covers between the branchial openings. 
Antero-laterally, between the foremost gill-opening and the 
jaw, they pass over the fore part of the branchial chamber. 
Here, as well as between the several branchial openings, they 
are continued upon the dorsum of the animal as far as the 
lateral septum, forming what may be called the ' cervicalie svr 
perfidalis superior * 

This superior muscle is quite as strongly marked as that 
on the under surface of the throat and branchial chamber. 
It may be described as arising on the dorsum of the animal 
and extending downwards to meet the inferior muscle. Taking 
this view of it, we find it to arise from the projecting upper and 
back part of the skull, from the anterior edge of the scapular 
part of the shoulder-girdle near the fin and, slightly, from the 



274 PBOFESSOB HUMPHBY. 

dorsal surface of the fin, where it represents the trapezius and 
meets the latissimua dorsi, also from the lateral septum be- 
tween the shoulder-girdle and the skull. In the last situation 
the fibres are traceable down, beneath the septum, to the 
lateral parts of the vertebrae and to the fibres of the^ deep 
portion of the ventral muscle pas.'^ing forwards to the skull. 
The fibres descend forwards and outwards, over the branchial 
chamber and the gill covers, and are continuous with the 
ascending fibres of the cervioalia superfidalis infeiHor. They 
pass over and are slightly connected with the branchial car- 
tilages in the same manner as the fibres of the last-named 
muscle are related to these, cartilages beneatL 

The points of difference between the superficial brachio-cephalic 
stratum in this animal and in Lepidosiren (p. 258) are slight, arid are 
' caused partly by the difference in the gill openings. The segmenta- 
tion from the deeper part of the stratum behind the girdle is less 
distinct in the Dog-fish ; and the stratum in front of the girdle does 
not, in this animcd, expand over the dorsal muscle as it does in 
Lepidosiren. There is further not that distinctly segmented deeper 
layer of the snperBcial stratum in or near the mesial line which I 
have called 'genio-hyoid' in Lepidosiren (p. 261). The cervuxdis super- 
fidalis in/erior I suppose to represent, as in Lepidosiren, the in- 
ferior part of the subcutaneus colliy the sterno-deido-mastoideus, the 
mylo-hyoidsuSj the anterior belly of the digastricus and the genio- 
hf/oideus; whUe the cervicalis auperficialis superior represents the 
superior part of the sv^cutaneus colliy the depressor mandibulcey the 
levator arcuum, the trapezius with the cervico-humerals and rhom- 
boids. 

Beneath the cervicalis auperficialia sup., which has been 
partly cut away in Fig. 28, is seen a strong muscle {Levator 
scapulas) arising from the lateral septum and from the sides 
of the vertebrae beneath it. The fibres pass backwards and 
downwards to be inserted into the under surface of the scapula 
where they meet the fibres of the lateral muscle, which I have 
called serratuSy coming forwards to the same cartilage. 

The division and reflection of the cervicalis superf. infi 
from the middle line, as represented, on the right side in 
Fig. 29, brings into view the continuation of the deeper layer 
of the ventral muscle, which may be called cervicalis profundus 
(C, p.\ from the fore part of the coracoid to the hyoid and 
the glossal cartilage. It constitutes a thick well-defined muscle, 



4 



MUSCLES OF THE SMOOTH DOO-FISH. 275 

like its homologae in Lepidosiren, and presents transverse 
septa corresponding in number and position with the branchial 
cartilages. At its origin this muscle, like the cerv, super/. , is 
connected with the coracoidal septum as well as with the 
coracoid ; and it extends with the septum for some little dis* 
** tance upon the ventral aspect of the fin. The proper muscle 
of the fin also derives origin from the septum as well as from 
the coracoid. The septum therefore brings into relation and 
gives origin to the cervicalis sup. inf. and the cervicalis prof, 
in front, and to the pectoraly ventraly and fin muscle behind, 
linking them all together and to the coracoid \ 

The fin-muscle upon each surface is, in consequence of the 
different direction of its fibres, like the muscle upon the median 
fins (see pp. 257, 280), in great measure segmented from the 
extensions of the lateral muscle which spread upon the fin. 

We have traced the superficial stratum of the ventral muscle 
(cerviccdis superfixyialis), over the shoulder*girdle and over the 
superficial branchial cartilages, and the deep stratum {cervicalis 
profundus) forwards from the coracoid to the hyoid. The 
deepest stratum of the ventral muscle passes from the deep 
( surface of the girdle upon the deep branchial cartilages*. Su- 

periorly, beside the bodies of the vertebras, it is continued 
from one cartilage to another, and from the uppermost cartilage 
to the base of the skull, forming a continuous series of alter- 
nating cartilages and muscles, like the ribs and the intercostals. 
Furthermore some of the fibres are reflected from the hindmost 
branchial cartilage upon the oesophagus, contributing to the 

^ The course of the nerves to the fin accords with the disposition of the 
mnscles, as compared with those of Lepidosiren and of higher animals. They aU 
pass behind the girdle, except one which penetrates the edge of the girdle imme- 
diately behind the point of articulation of the fin. They traverse the ventral 
muscle in a line behind this point, having above them the part of the ventral 
muscle attached to the scapula which I have designated «erratiz«, and having be- 
neath them the part of the ventral muscle attached to the coracoid, and forming 
a deeper stratum than the pectoral, which represents the costo-eoracoid. 

' The deep branchial cartilages, five in number, commence above, that is 
beneath the vertebral bodies, by flattened, slightly curved ends, in dose contact 
with the mucous membrane of the pharynx. At the level of the sides of the 
bodies of the vertebrse they suddenly thicken and turn downwards, running round, 
in contact with the pharynx, to the median line, where they coalesce with those 
of the opposite side. The superficial branchial cartilages are nearly subcutaneous, 
% separated from the skin by the fibres of the superficial cervical muscles only. At 

their upper or vertebral ends they are connected by fibrous tissue with the deep 
cartilages ; and their lower ends expand and underlie the deep cartilages which 
are also expanded. 



276 PBOFESSOR HUMPHRY. 

muscular coat of that tube and forming a diaphragm between 
the branchial and pericardial chambers. Some of the fibres 
also pass from the oesophagus to the vertebral column. 

The muscular coat of the oesophagus is further augmented 
by a small muscle on either side which extends, from the base 
of the skull and the foremost vertebra, upon the upper flat 
ends of the branchial cartilages and upon the oesophagus. 

A second or post-cardiac diaphragm is formed by a layer 
of the fascia transversalis reflected from the hinder edge of 
the girdle upon the oesophagus. It is not accompanied by any 
muscular fibres. 

Ventral Fin (Fig. 30). The dorsal muscle (Gl) of the 
ventral fin is broad and square-shaped^ and in part continuous 
with the superficial strata of the ventral muscle, whereas, 
behind, it overlies the ventral muscle and is easily dissected 
from it. It thus affords a good example of transition from 
continuity with the lateral muscle to superposition with regard 
to it. As it descends upon the fin it blends with the proper 
muscles of the fin and extends also upon the clasper. It repre- 
sents the gluteus muximtis, together, probably, with the sariorius 
and tensor voffince femoris (see also Ceratodus, p. 283). I do not 
trace any of the fibres of the caudal part of the rentral muscle 
into the back of this fin-muscle or into the back of the fin 
or the clasper. The caudal miiscle is, however, largely inserted 
into the pelvis behind, as is the ventral abdominal, muscle 
in front. Near the mesial line the pelvis quite interrupts the 
ventral muscle, cutting oflF the caudal from the abdominal part, 
and lying in contact with the visceral cavity. More laterally, 
however, the pelvic cartilage is imbedded in the ventral muscle, 
the fibres of which are continued, on both its superficial and 
deep surfaces, from the caudal to the abdominal region. 

The following description of three transverse sections of 
the animal, together with the drawings (Figs. 31, 32 and 33) 
and the accounts of them at p. 278, shew the relations of the 
lateral muscles to the septa, to the vertebrae and their processes 
and to the median fins. 

A transverse section of the animal through the anterior 



MUSCLES OF THE SMOOTH BOG-FISH. 277 

dorsal fin (Fig. 31) shows the lateral septum {L.8.) passing 
from the laterally projecting margins of the body of the 
vertebra, quite on a level with its lower surface which is flat- 
tened or slightly concave. It first takes a sweep downwards, 
then ascends to the part where it appears on the surface. 

Beneath it, the ventral muscle is extended over the abdo- 
minal cavity and presents one thick oblique septum (C) indi* 
eating the commencement of the medio-ventral part of the 
muscle {M. F.). Above the lateral septum septa radiate from 
the middle of the neural arch about the position of the articu- 
lating processes. There is no neural spine, or a very short one. 
The median neural septum is continued, from the summit of 
the neural arch, to the cartilage of the fin, and there splits 
into the septa {A. A.) which separate the fin-muscles, on the 
two sides, from the lateral muscles. These last-mentioned 
septa resemble the other septa of the lateral muscle, though 
they are rather thicker; and their relations indicate the fin- 
muscle to be a segment of the lateral muscle. The cartilagi- 
nous fin-ray extends far beyond the fin^muscle into the fins ; 
and its extremity is embraced by the homy subcutaneous fibres 
of the fin, which pass on either side of it 

In a section (Fig. 32) through the posterior dorsal fin and 
the anal fin the lateral septum is seen passing from the side 
of the haemal canal, at a point below the body of the vertebra, 
about corresponding with the point, above, from which a septum 
passes from the side of the neural canal, a point, that is, answer- 
ing to the articulating processes. The neural spine is some- 
what pronounced, as is also the haemal. The cartilaginous rays 
extend into the haemal ajs well as into the dorsal fin. 

In a section (Fig. 33) through the caudal fin the relations of 
the lateral septum to the hiemal canal correspond with those in 
the section just described. The neural and haemal spines, 
especially the latter, are much more prolonged, extending 
quite to the bases of the respective fins; and the homy fila- 
ments of the fins embrace their ends^ much as, in the dorsal fin, 
they embrace the free ends of the cartilaginous rays. These 
rays are here absent, and there are no fin-muscles. 

Thus the neural and haemal spines are reciprocal with re- 
gard to the cartilaginous rays. When these are developed 



278 PROF. HUMPHRY. MUSCLES OF THE SMOOTH DOG-FISH. 

and penetrate between the lateral muscles the spines are short 
or abortive ; and, on the contrary, when the rays are abortive, 
the spines run on into the fins. 



DESCRIPTION OF FIGURES. (PI. XIII.) 

Fig. 28. Dissection of the uppet sur&ce of the fore part of the 
Dog-fish, to show the angular prolongations of the septa of (if. D.) 
the medio-dorsal part of ^e lateral muscle. Z. D., the latero-dorsal 
part of the same. L. V., the latero-ventral, and M. F., the medio- 
yentral. Zot d.j latissimus doni, which has been partly reflected 
from the fin-muscle. P, pectoral. C. 8. s., cervicalis superficialis 
superior, its supra-branchial portion. 2>. F., the dorsal fiu. 

Fig. 29. Dissection of the under surface of the fore part of the 
Dog-fish. M, v., the medio-ventral portion of the lateral muscle. 
Py its superficial, or pectoralis, layer advancing to the fin and to 
{C. S.) the coracoidal septum. C. 8, i,, the cervicalis superficialis 
inferior, which, on the left side of the animal, is in situ, but, on the 
right side, has been reflected to shew (C. pr,) the cervicalis profundus, 
and the superficial branchial cartilages. The hinder fibres of the 
cervicalis pr. are seen passing upon the fin and connected with the 
fiu-muscle. G. II., the glosso-hyaL C, H,, cerato-hyaL B, bran- 
chiostegal rays. 

Fig. 30. Side view of a portion of the hinder part of the Dog-fish 
to show the relation of the dorsal muscle (GL) of the ventral fin (F. F.) 
to (Z) the lateral muscle. P, D. F.^ the posterior dorsal fin. CL CI., 
the claspers. 

Fig. 31. Section through the anterior dorsal fin and the body 
wall. L. S., lateral septum. A, septum between the dorso-latersd 
muscle and the fin-muscle, passing to the root of one of the cartila- 
ginous fin-rays, and continued on, as a median septum, to the dorsal 
spinous process. A\ similar septum to the fin-ray above. B, septum 
from neural arch separating Z>. Z., the dorso-lateral muscle from M, Z>., 
the medio-dorsal muscle. (7, septum between (F. Z.) the ventro- 
lateral and (M. V.) the medio*ventral muscle. The parts of the 
vertebra which are ossified are distinguished from the cartilaginous 
parts by not being dotted. 

Fig. 32. Section through the posidorsal (P. d, F.) and the anal 
(A, F,) fins. r, the ventral muscle behind the part where it is 
divided into medio-ventral and ventro-lateral. 2>, septum between 
the ventral muscle and the anal fin and fin-muscle. A, septum 
between the dorsal muscle and the dorsal fin and fin-muscle. 

Fig. 33. Section through the caudal fin showing the prolongation 
of the neural and hiemal spines into the fin. Z. S., lateral septum. 
N. S. is the divided end of ^q overhanging neural spine; and ZT. S, 
is the divided end of an underlying haemal spine. 



^ 
V 



THE MUSCLES OF CERATODUS. By Professor 
Humphry. (Plate XIV. Figs. 34 to 38*). 

Lateral and Mesial Fin-Muscles. 

The lateral muscle presents the usual general piscine charac- 
ters. There is, however, none of the superficial, soft, coloured 
stratum often found over and near the lateral septum'. The 
ventral intermuscular septa simply slant from the lateral septum 
backwards and downwards to the ventral line* Traced inwards 
they pass obliquely forwards to the ribs, or to the transversalis 
fascia in the trunk, and in the tail to the haemal spines and 
the median septum. Near the pectoral girdle the direction of 
their slant is reversed. The dorsal intermuscular strata, at first, 
slant backwards and upwards from the lateral septum ; then 
they curve forwards to the line at which the dorsal fin-muscles 
impinge upon the lateral muscle. Here the direction is again 
suddenly changed ; and they slant backwards and upwards, 
through the fin-muscle, with a greater obliquity than they 
present near the lateral septum. The septa in the muscle 
extending upon the side of the dorsal median fin are distinctly 
continuations of the septa of the lateral muscle, though their 
direction is somewhat different. The same disposition is ob- 
served throughout the tail ; except that as they are traced back- 
wards the curves of the septa disappear; and in the hinder part of 
the tail the septa extend as straight lines, obliquely backwards 
and upwards, through the dorsal part of the lateral muscle and 
onwards, with merely an increase of obliquity, through the fin- 
muscle. In like manner the septa of the ventral part of the 
lateral muscle, behind the anus, merely acquire an increased 
slant backwards when they traverse the muscle upon the sides 
of the median fin. 

Near the lateral septum the fibres of the lateral muscle, 
ventral and dorsal, take an antero-posterior direction, a direc- 

^ For the opportnnitj of dissecting this animal I am indebted to the kind- 
ness of Prof. Flower, of the Boyal College of Surgeons. It was dissected for me 
by Mr Anningson. The anatomy of Ceratodos, with the exception of the moscles, 
has been foSy described by Dr Giinther, in an admirable paper contained in 
the Phil. Tram, of the Boy. Soc. for 1871. 

* This was also absent in Lepidosiren and Ceratodtw. See obs. p. 294. 



280 PROFESSOR HUMPHRY. 

tion that is parallel with the direction of the septum and of the 
axis of the animal. As they become more distant from the 
septum they acquire a slant ; t]^e fibres of the ventral muscle 
inclining a little downwards and backwards and those of the 
dorsal muscle a little upwards and backwards. Near the ventral 
mesial line the direction of the fibres is again antero-posterior ; 
but near the dorsal mesial line the fibres still retain a slight 
obliquity. Dissected from without inwards the fibres of the 
deeper parts of the ventral muscle are found to preserve much, 
the same direction as do those near the exterior; that is, no 
difference is traceable which would indicate an incipient divi- 
sion into external and internal oblique, such as was observed in 
Lepidosiren (p. 256) and more markedly in Cryptobranch 
(p. 11). We notice that as they approach the interior they 
become intermingled with an increasing number of threads 
of fibrous tissue which take the place of the muscular fibres. 
In some places a tolerably distinct stratum of these, pass- 
ing from rib to rib or septum to septum, lies upon the trans- 
versalis fascia. There are no muscular fibres beneath • the 
level of the ribs to represent a ti-ansversalis muscle. This de- 
creasing proportion of muscular fibres towards the interior of 
the abdominal wall is interesting as an indication that the 
development of its muscular tissue become less active as we 
approach the interior (see obs. on the Tadpole, p. 301). 

The intermuscular septal planes being continued into the 
muscle of the median fin, it follows that the muscular planes aire 
likewise so continued, and that the fin-muscle is an extension of 
the lateral muscle. There are, however, some distinguishing 
points. First, the direction of the fibres is different. The 
fibres of the fin-muscle are nearly parallel with the fin-rays, 
that is, have a much more vertical course than those of the 
dorsal muscle ; and the change in direction takes place suddenly, 
the fibres of the fin-musde impinging upon those of the dorsal 
muscle at an angle which approaches to a right angle. Hence 
the one set are segmented from the other ; and a layer of the 
superficial fascia, as described in Lepidosiren, and shewn in 
Fig. 31 of Dog-fish, passes between them, covers the edge of the 
lateral muscle, and meets its fellow in the mesial line, or comes 
into relation with the fin bones and the vertebral spines ; while 



MUSCLES OF THE CERATODUS. 281 

the other, or outer, layer Is continued upon the surface of the 
fin-muscle and the fin. The fin-muscle and fin are continued in a 
channel between the two fascial layers just mentioned, and 
are transversely segmented by septa crossing the channel and 
continuous with the transverse septa of the lateral muscle. The 
fin-ray bones are ossifications in the meeting-points of the septa 
of the two sides in the median line; just as the spinous 
processes and ribs are ossifications in the median or other parts 
of the transverse septa of the lateral muscle. The fin-muscle is 
accordingly divided into bundles, which correspond in num- 
ber with the interspinous and fin-ray bones ; and each bundle 
is inclosed in a fascial sheath which separates it from the 
adjacent bundles as well as from the lateral muscle. Each 
bundle also is composed of fibres running parallel with the fin- 
ray and springing from the surrounding sheath, chiefly from the 
basal part of the sheath which rests upon the lateral muscle, 
but partly from the septa which separate them from the fibres 
of contiguous bundles. 

The superficial or exterkal FASCII is unusually thick. It 
is connected by areolar tissue with the derma on the one side- 
face, and with the septa of the lateral muscle on the other ; 
these connecting it, as in Lepidosiren and Cryptobranch, with the 
fibres of the lateral muscle and with the deep, or internal, or 
transversalis fascia. It is also continuous with the transversalis 
fascia along the ventral edge of the animal ; and, in the abdo- 
minal region, the blended fasdsB of the two sides are united in 
the median line, forming a median septum. Behind the pelvis 
the blended superficial and deep fasciae on the one side are 
separated from those of the other by the bowel passing between 
them to the anus ; and a sheath formed by them is continued 
upon it (see also Fig. 23, 0). 



Muscles of Pelvic and Pectoral Fins. 

The cartilaginous pelvis, with its smooth anterior, or pre- 
pubic, styliform cartilage and its lateral comua, resembles that 
of Lepidosiren (p. 257 and Fig. 23) ; but the comua are shorter, 
and each is forked (Fig. 34 0), terminating in two blunt prongs. 



282 PBOFSSSOR HUMPHRY. 

into the hinder of which two intermuscular septa converging 
are inserted. Behind, the pelvis terminates in a semilunar 
edge, with a condyloid projection on either side for the articu- 
lation of the paddle-like fin\ 

The styliform process is enclosed in a sheath of the hlended 
fasciae, as in Lepidosiren. The comua lie in the lateral muscle, 
the fibres of which are attached to their anterior and posterior 
surfaces. The under surface of the pelvis is covered, on each 
side, by a thick muscular mass, the fibres of which are parallel 
with those of the lateral muscle, though they are separated from 
them by a cleft This mass is divided into two nearly equal 
portions — ^a median or 'plantar' and a lateral or 'dorsal;' and the 
line of division between them corresponds with, and is a conti- 
nuation of, the line of the inferior edge of the fin. The dorsal 
or lateral portion {GL Fig. 34s) passes upon the dorsal or outer 
surface of the fin ; and the plantar portion passes upon the ven- 
tral or inner surface of the fin nearly to its extremity*. Each, 
in its course upon the pelvis and upon the fin, is marked by 
transverse inscriptions, like those of the caudal lateral muscle. 
These inscriptions are transverse upon the proximal thick part 
of the fin ; but, on the more distal part of the fin, they be- 
come oblique, slanting firom the middle or axial line of the fin, 
backwards and outwards towards the margin of the fin. They 
correspond with and overlie the lines of division of the axial 
cartilage of the fin into pieces; and they extend over the rays 
which pass, in a penniform manner, from those lines. They do 
not extend between the pieces of the axial cartilage so far as 
the margin of the fin, but only about half-way from the axis to 
the margin. In the intervals between them the muscular fibres 
are directed longitudinally, parallel, that is, with the axis of the 
fin. In the more marginal parts of the fin the muscular fibres 
slant off obliquely, from the middle longitudinal set, which form 
the proper muscle of the fin, towards the margin; and the two 
sets are to some extent segmented from each other, in the same 

1 Gtniher, I. e, p. 685 and fig^nres. In C^iobraneh the prepabio cartilage 
is not oontinuoos with the pelvic cartilage, as in CeratodoB and Lepidosiren, but 
jointed to it by fibrous tissne. It is also less cylindrical and smidl, and bifur- 
cates anteriorly into two flat unqymmetrical boms. 

' I use the words 'plantar' and * dorsal* in reference to those portions of the 
fi^n-muBcle, to indicate their correspondence with the respective groups of 
musdes passing to the hind limb in lugher animals. 



MUSCLES OF THE CERATODUS. 283 

manner ajs the muscles of the median fin of the body and tail 
are segmented from the adjacent parts of the lateral muscle, but 
less distinctly. Towards the extremity of the fin the distinction 
between the two sets becomes less marked; and, at last, the 
middle set of fibres are continued without interruption to the 
edge of the fin (Fig. 34). This similarity in the construction 
of this almost uniquely bipinnate fin or paddle and in the 
arrangement of its muscles, to the construction of the tail and to 
the arrangement of the muscles of the tail, is highly interesting. 
It is strongly confirmatory of the view I have formed, from the 
dissection of this animal and Lepidosiren, that the muscles of 
the mesial fin are derivations from the lateral muscles, present- 
ing varying degrees- of continuity with, or segmentation and 
separation from, them. 

The muscle on the dorsal surface of the fin is not derived 
entirely from the girdle. It receives a direct accession from 
the lateral muscle in the form of fibres which converge, from 
several of the segments of the lateral muscle, upon the fin, and 
which blend in one of the septa of the fin muscle (Fig. 34, Q'L). 
Of these fibres the foremost, running parallel or nearly so with 
those of the lateral muscle, are continued uninterruptedly into 
them. The hinder fibres, crossing that muscle more at an 
angle, are segmented from it, lie upon its surface, and are with- 
out difficulty dissected from it, affording, like the corresponding 
muscle in the Dog-fish (p. 276, Fig. 30), another illustration 
of gradual segmentation in consequence of increasing difference 
of direction. 

The accession to the dorsal fin-muscle represents not uncertainly 
the sheet of the ventral muscle extending upon the dorsal aspect of 
the corresponding limb in higher animals, which may be partially 
or wholly interrupted by the ilium growing through it, and which 
becomes more or less segmented into sartoriics, tensor vaginas femoria^ 
and glvJUuB, The part of the muscle derived from the pelvis repre- 
sents the deeper glutei, <&c. 

In like manner the muscle on the ventral aspect of the fin 
is not derived altogether from the plantar and internal or mesial 
portion of the sub-pelvic mass just described. It is joined by 
bundles of fibres from the deep surface of the hinder or ischi* 
atic part of the pelvis; and, moreover, some of the fibres from 



284 PROFESSOR HTJMPHRT. 

the lower edge of the ventral muscle passing forwards from the 
tail {Pt. Fig. 36) are reflected into it. 

This accession to the plantar fin-muscle from the caudal part of 
the ventral miiscle may be regarded as representing the caudo-appen- 
dicular seiies of Urodelans which are variously segmented into caudo- 
pedal, caudo-crural and caado>femoraL 

The disposition of muscles upon the pectoral fin, which is 
also a bipinnate paddle, resembles closely that of the muscles 
upon the ventral fin, except that they are formed altogether by 
fibres passing from the girdle, there being no accessions from 
the lateral muscle, no correspondents, that is, with the latissi- 
mus dorsi and pectoroMs, The fibres that arise from the sca- 
pular part of the girdle — ^the part above the joint with the fin — 
form a ' scapulo-brachial' muscle, which passes upon the dorsal 
aspect of the fin; and the fibres that arise from the coracoid 
part of the girdle — the part beneath the joint with the fin — 
form a ' coraco-brachial' muscle which passes upon the ventiul 
aspect of the fin. 



Brachio-Cephalic Muscles. 

Traced forwards the ventral muscle does not give off, sus in Le- 
pidosiren, a superficial brachio-cephalic muscular stratum ; though 
the superficial fascia, which represents it, is continued forwaixls 
over the gii'dle to the jaw. A representative of the cervical or 
cephalic portion {certricalis superficialia) of this muscular stratum 
is, however, found in front of the pectoral limb-girdle in the 
form of a muscular layer composed of fibres or bundles passing 
transversely, or in a slightly curvilinear manner, over the gill- 
cover (Fig. 34, C. «.). Traced upwards these fibres do not reach 
the lateral septum, but are attached to the girdle (the scapular 
part of it) above the fin, to the supra-scapula, and the back- 
wardly projecting cartilaginous occipital angle of the cranium. 
Anteriorly, they encounter and are attached to the opercular 
and sub-opercular bones, the hinder ends of which project into 
and interrupt the layer, and to the lower jaw. Inferiorly, they 
extend beneath the throat, and are lost in the extension of the 
superficial fascia. By this means only they acquire connection 



MUSCLES OF THE CERATODUS. 285 

with the rest of the ventral muscle. They take chiefly a trans- 
verse direction, and bear close resemblance to the corresponding 
cervicalis swperficialis of Lepidosiren and Dog-fish. The re- 
flection of it, as in Fig. 37, shews it to be inserted strongly into 
the cerato-hyoid cartilage {G. Ed)y also into the jaw near its 
angle, as well as into the opercular bones, and into the body 
of the jaw (Jf) between the angle and the symphysis. There 
are not any fibres running forwards from the hyoid to the 
symphysis and corresponding to those which in Lepidosiren 
(p. 261) I have supposed to represent the anterior belly of the 
digastric. 

This muscle is mentioned, and partly described by Giinther, p. 525. 
Lying beneath the outer or posterior half of the suboperculumy 
closely united with and projecting beyond it, so coming into relation 
with the fibres of the muscle just described, is a remarkable plate of 
cartilage, represented at S. o. c. Fig. 37, which is not mentioned by 
Giinther. It is quite separate from all the cartilages of the skull, 
and is connected only with the suboperculum, though lying near the 
hinder edge of the cei*ato-hyoid cartilage. There is not any similar 
cartilage under the operculum ; for the small cartilage described as 
pre-operculum by Giinther is quite separate from the opeix)ulum. 

Beneath the cervvoalis superfidalis, near the middle line, is 
the muscle which I have described in Lepidosiren (p. 259, and 
Dog-fish, p. 273), as the deep layer of the cervicalis superficialis. 
In this animal it is more separate from the remainder of the 
cervicalia sup. than it is in Lepidosiren; and it constitutes a 
distinct genio-coracoid (G. c. Fig. 37). It is flat, riband-like, 
and attached behind to half-an-inch of the anterior edge of the 
coracoid cartilage, and of the epicoracoid-bone (p. 294), near the 
middle line. It extends, forwards, to the under-surface of the 
hinder edge of the mandible near the symphysis. It passes 
beneath the hyoid, and is not connected with it ; still its fore- 
part may, and probably does, represent the genio-hyoid of 
other animals or the hyo-mental (p. 325). 

The DEEP LAYER OF THE BBACHIO-CEPHALIC EXTENSION is 

disposed much as in Lepidosiren. It first, in its lateral part, 
encounters the peculiar bone, or first rib, which appears to be 
formed in the lateral part of one of its septa. From the ante- 
rior and outer surface of this bone is given off a thick muscle 

VOL. VL 20 



286 PROFESSOB HUHPHBY. 

(serratua), which is a continuation of this stratumi to the under- 
surface of the scapula. A little more anteriorly and near the 
middle line the brachio-cephalic is inserted into the coracoid 
cartilage and the epicoracoid septum which quite interrupt it. 
In front of the coracoid it is continued as a thick muscle, but 
without septa, to the hyoid forming a coraco-hyoid, the repre- 
sentative apparently of the stemo-hyoid. The middle part is 
inserted into the base of the cone-shaped glosso-hyal by a flat 
tendon which passes beneath the basi-hyal. The fibres of its 
lateral part, which are situated in rather a deeper level, are 
inserted into the basi-hyaL 



DESCRIPTION OP THE FIGURES OF CERATODUS 

(PI. XIV.). 

Fig. 34. A side-view of the animal, shewing D, the dorsal, and 
V, the ventral muscle^ and the lateral septum between them. 

P, the styliform prepubic cartilage turned out of its median 
sheath, and inclined upon the left side of the TentraJ muscle. C, 
comua of pelvic cartilaga Gl., dorsal-fin muscle passing from the 
pelvis, upon the dorsal surface of the ventral fin. Gl/, accession to 
dorsal-fin muscle from lateral muscle. Ay the anus. S, F., the 
superficial fascia reflected from the outer-surface of the ventral 
muscle. It joins {T) the transversalis fascia coming from the inner- 
surface of the ventral muscle. The two united are continued upon 
the boweL 0., opercular bone. iS, 0., sub-opercular bone. C, 8, «., 
cervicalis superficialis superior muscle (its supra-branchial portion) 
passing over gill-cover. CBr., coraco-brachial mass passing upon 
the ventral aspect of the fin. S. JBr., scapulo-brachial muscle passing 
upon the dorsal aspect of the fin. 

Fig. 35. A piece of the lateral muscle with one of the myotomes 
dissected out to shew the sclerotome, or intermuscular septum, /. if., 
passing inwards and forwards above and below the lateral septum to 
jP, the fin-ray bone, and Ef the rib, in which it terminates, as well 
as in the side of the body of the vertebra between them. ^, a 
rib in front. The transversalis fascia is seen between the two ribs 
with an outer coating of tendinous fibres which slant, from above 
downwards (ventrally) and forwards, in a direction that is difiPerent 
from those of the muscular fibres above them, and corresponding with 
that usual in internal intercostals or depressores costarum. The cat 
ends of the fibres of the ventral muscular bundles are seen in front 
of the gap; whereas the doi-sal fibres have been cleared away in the 
corresponding situation, and the under-sur£BM;e of the septum next 
ac^aoent haa been there exposed* 



MUS0LB8 OF THE CEEITODUS. 287 

Fig. 36. The pelvis fmd fin with part of the lat^^l muscle. 
P, the prepubic cartilage. GL, the dorsal muscle from pelvis to fiu, 
and Gl, , the accession to it from the ventral muscle. The latter has 
been divided and reflected to shew PL, the plantar muscle from the 
pelvis to the fin, and Fl.\ the accession to it from the caudal part of 
the ventral muscle. 

Fig. 37. Dissection of the deeper muscles in the under-surface 
of the throat. F, the ventral, or brachio-cephaJic part of the ventral 
muscle. S, F., superficial fascia passing over (7, the coracoid to 
Sj the symphysis of the mandible. C»8.s., cervicalis auperfioialis 
superior inserted into, S, o., the sub-operculum. S, o,c,y the sub- 
opercular cartilage. C, A., the cerato-hyal, and M, the mandible. 
C. jET., the coraco-hyoid inserted into (r. A., the glosso hyal, and 

B. A., the basi-hyal. G. c, the genio-coracoid of the right side, (r.c, 
the genio-coracoid of the left side turned back in its middle and 
hinder part. C. Br., coraco-brachial. 

Fig. 38 shews the connection of the fore part of the ventral 
or brachio-cephalic muscular mass, and its septa with the coracoid. 

C. C, the coracoid cartilage. (7', ^e epicoracoid bone. The muscular 
fibres have been left near the. middle line, but dissected away more 
laterally to shew the septa which here slant, from the surface, back- 
wards and inwards. The foremost septum is continued into the 
coracoid. The next has the peculiar rib-bone, J?, projecting into it. 
It is continued into that bone. (7. Br,, the coraco-brachiid muscle 
extending from the girdle, beneath the joint with the fin, upon the 
venti*al surface of the fin. S. Br.^ scapulo^brachial muscle extending 
from the girdle, above the joint with the fin, upon the dorsal surface 
of the fin. 



NOTES ON THE MUSCLES OF THE GLASS-SNAKE. 
Pseudopua PaUasii \ By Professor Humphry. 

The hind-limb is a mere slight scaly projection from the margin 
of the ventral articular plate, where the latter is being rounded 
oflFto form the anterior wall of the cloaca. The pelvic girdle is 
attached, above, to the transverse process of one vertebra". 

1 This animal was diseeoted for me by Mr Aimiagson. 

' The ilium extends a little above the transverse process; and the inner 
side of the ilium is applied upon the extremity of the transverse process (the 
second behind the last rib) ; a ball and socket is formed, the round end of the 
transverse process being reoeived in a shallow socket in the ilium ; and there is 
a synovial cavity with a thick surrounding capsule. The arrangement permits 
the lower ends of the two pelvic bones to swing backwards and forwards upon an 
axis traversing the transverse processes of &e two aides. This movement is 

20—2 



288 PHOFESSOB HUMPHRY. 

Beneath, it is free in the abdominal wall, i. e., not connected 
with that of the opposite side. The small osseous limb marks, 
by its point of connection with the pelvis, the division between 
the iliac and the pubischiatic, or ischiatic, parts of the pelvis. 

The rectus abdominis is a more distinct and continuous 
sheath than in the Snakes I have seen. It is joined by the 
successive bundles of the ohliquua extemus abdominis passing 
from the ribs into its upper or abdominal surface. Traced back 
it is quite interrupted by, or inserted into, the pubischiatic 
bone; and the hindmost division, or bundle of the external 
oblique muscle is inserted with it, and above it, i.e. deeper 
than it, into the same bone. 

Examined more carefully the rectiLS abdominis {longissimtis 
abdominis of Ftirbringer*) is found to be composed, first, of 
bundles directed, forwards and outwards into it, from the 
extremity of the ribs and the costal cartilages, the hindmost 
of the bundles being attached to the mesial part of the pelvic 
bone (they resemble detached portions of external intercostal 
muscles) ; secondly, of bundles passing backwards and down- 
wards, into it from the ribs nearer the vertebrae. These two 
sets of bundles lie in the same oblique lines; and they are 
confluent in the substance of the rectus. That muscle may 
therefore be said to be formed, either of bundles passing from 
• the ribs near the vertebrae into it, travelling for a distance in it, 
and then leaving it to be attached to the free ends of the 
libs and the costal cartilages ; or, it may be described as being 
formed of fibres which take the opposite direction, that is, 
arising from the free ends of the ribs, joining it, and then 
leaving it to be inserted into the ribs near the vertebrae". 

These bundles run chiefly into the deeper surface of the 



restricted by a fibrous band conneoting the ilium with the transyene process 
fiext in front of it, the transverse process that is next behind the last rib. In 
an articulated skeleton in the Cambridge Mnseum the ilinm is connected 
directly with the transverse processes of two vertebra. 

The limb consists of one longer bone next the pelvis, carrying a smaUer 
bone, a mere nodnle, upon its end. These bones should scarcely be named 
tibia and femur. They are as much representatives of the other Umb-bones as 
of these two. 

^ Die Knoeken und Mutkeln der Extremitaten der ichlangen&hnliehenSauriem, 
von Max Fiirbxinger. Leipzig, 1870. 

* This is the case also in Snakes; though in them the bundles are finer and 
run more into the seiites, and form a less distinct antero-posterior muscle. 



MUSCLES OF THE GLASS-SNAKE. 289 

IcUerai part of the rectus. Its more mesial part is formed, 
wholly or nearly, of antero-posteriorly directed fibres*. 

Behind the pubischiatic bone is a continuation of the series 
of external oblique muscular fibres, backwards, to the cloaca 
and into the caudal muscle on the side of the cloaca^ so con- 
stituting an ischio'cloacalis and an ischio-coccygeus. These, 
it will be understood, are separated from the rectus and the 
ohliquus by the pelvic bone. 

Badiating forwards and inwards to the middle line from 
the median extremity of the pubischiatic bone is a pyramidaUa. 
It is quite distinct, though thin ; and it is broad. It lies along 
the inner side of the rectus ; and the pyramidcUes of the two 
sides occupy the space on the ventral surface of the abdo- 
men which is left uncovered by the recti diverging as they 
pass backwards to the pubischiatic bones*. This muscle is not 
described by Furbringer. 

The gracilis passes from the pubischiatic bone, where it is 
partially continuous with the rectus abdominis, into the inner 
or plantar side of the limb, and is attached to both the bones 
of the limb. Beneath the gracilis some deeper fibres, passing 
from the pubischiatic bone to the upper limb-bone, represent 
the adductors. 

The deeper stratum of the abdominal muscle, the stratum, 
that is, beneath the level of the ohliquus extemvs and the 
rectus, which extends more laterally than they, and in which 
the ribs are situated, and which therefore corresponds with the 
obliquus intemus, if traced backwards is found to pass, in great 
measure, superficial to the ilium, into the ventral caudal muscle. 
Some fibres are, however, inserted into the ilium ; and a tole- 
rably distinct part of it passes, from the hindmost rib, to the 
ilium, constituting a quadratus lumborum and representing 
serially the intercostals. 

If this internal oblique stratum is traced forwards, 
from the tail, a larger number of its fibres are found to be 
inserted into the hinder edge of the ilium, constituting an 
iliO'Cavdal; and one distinct bundle of fibres is found to pass 

^ These might be described fts a third set. They are, howerer, probably 
oontinuons indirectly with the others. 

> See similar disposition in Pteropus, Joum, AnaU 



290 PROFESSOR HUMPHRT. 

beneath the ilium and to terminate in a tendon which runs on 
into the abdominal muscles ; so that though the ilium lies deep 
it does not lie quite so deep as it does in Snakes, the ventral 
muscle being partially inserted into it and even partially 
travelling beneath it. 

When the layer of the abdominal muscle travelling over the 
ilium is divided, a muscle is seen passing from the outer sur- 
face of the ilium upon the upper limb-bone. It is a repre- 
sentative of the deep glutei. 

We do not discover any part sufficiently distinct to deserve 
the name rectus fenvoria ; though such a muscle is described by 
Fiirbringer. 

The transversalie stratum of the abdominal muscles is repre- 
sented by a sheet of muscular fibres passing from the sides of 
the bodies of the vertebrae, forwards and laterally, upon the 
imder surface of the ribs, so forming depressores costarum. 
Some of these are traceable beneath three or four of the ribs. 
They decrease in thickness as they extend laterally ; and they 
are scarcely traceable beneath the mesial parts of the ribs. The 
nerves lie between them, on the one side, and the ribs and the 
more superficial strata, on the other side. There is no sub- 
vertebral rectus. The rectus capitis is strong, arising from the 
anterior five or six vertebrse, also from the side of the haemal 
spines of the three foremost vertebrae, and is inserted into the 
lateral part of the occipital. It encroaches upon the region 
usually occupied by the longus colli. These two muscles are 
however, as before shewn (p. 17), segments from one sub- 
vertebral mass« 

Brachio-Cephalic Muscles. 

Near the girdle the components of the rectus abdominis muscle 
radiate and separate into planes. The most superficial passes 
over the sternum and girdle, beneath the throat, over the 
lower jaw and side of the neck, and is lost in the tissue upon 
the back of the neck and head, thus constituting the sub- 
cutaneus colli. 

Next beneath this, and crossing the direction of its fibres, 
is a tolerably defined muscle, the depressor mandibtdce (cervid 
submaxiUai^ of Fiirbringer), arising firom the fibrous tissue 



1> 



MUSCLES OF THE GLASS-SNAKE. 291 

over the cervical spines and running, forwards and downwards, 
to the side of the angle of the lower jaw. 

A little in front of the clavicle these two muscles are blended 
in a transverse inscription about two lines in length. This is 
supposed to be a representative of the acromion by Rudinger. 
It extends also through the subjacent stratum. 

The next plane, partially interrupted by attachment to the 
sternum and clavicle, is continued forwards. Its more mesial 
portion runs under the clavicle, beneath the throat, to the hyoid 
and lower jaw as superficial stemo-hyoid (Fiirbringer) and gmio- 
hyoid, hyo-mental, and mylo-hyoid. The more lateral part — eter- 
nO'Cleido-niastoid — ^a flat band slightly connected on its deeper- 
surface with the clavicle \ runs to the back of the suspensorium 
(squamous or mastoid). It is superficially interrupted or di- 
vided into two— an anterior and a posterior part — ^by the in- 
scription above mentioned. 

More laterally still, that is, above the sterno-mastoid and 
rather on a deeper level than it, arising from the anterior edge 
of the upper, and scapular part of the clavicle, and on the 
same level with fibres of the stemo-hyoid and continuous 
with it, is a muscle which is called by Fiirbringer a part of 
the sterno-mastoid, but which appears to me more properly 
to be called trapezius. It passes forwards to be inserted into 
the back of the suspensorium, immediately behind the sterno- 
mastoid, with which it is more closely connected near the skull 
than it is near the girdle. 

These constitute the SUPERFICL^ BRACHIO-CEPHALIC 
STRATUM; those next described constituting the DEEP 

BRACHIO-CEPHALIC STRATUM. 

The third, or deepest plane of the ventral muscle passing to 
the girdle, consists, first, of a broad muscle passing from the 
first rib, in nearly its whole length, to the hinder edge of the 
deeper surface of the scapula ; secondly, of a broad bundle from 
the mesial part of the second rib to the hinder edge of the 

^ It Ib rather remarkable that the clavicle in its upper or Bcapnlar part is 
in, that is interrupts, this plane. Whereas in its lower or stemid part it lies 
beneath it, and interrupts the deepest plane of the brachio-oephalio mass. The 
superficial stemo-hyoid muscle passes quite superficially with regard to the 
clayicle, with the exception of a few of its lateral fibres which are continuous 
with the trapezius, and which are attached to the clavicle ; whereas the deep 
stemo-hyoid is interrupted by the clavicle. 



292 PROF. HUMPHRY. MUSCLES OF THE GLASS-SNAKE. 

scapula ; thirdly, of a narrower longer bundle from the mesial 
part of the third rib to the hinder edge of the coracoid near 
the sternum and to the sternum. It passes on also over the 
coracoid to the clavicle. Fourthly^ a long thin band is attached 
to the coracoid. It runs nearly straight backwards over about 
a dozen ribs and, finally, blends with the intercostals. Of these, 
the first two appear to belong to the 'serratus' group, and the 
last two to the * costo-coracoid * group. The absence of a limb 
precludes the opportunity of ascertaining whether the nerves 
to the limb pass, as they usually do, between the two groups \ 

The levator scapulce is a continuation of this plane onwards 
from the anterior edge of the upper part of the scapula, beneath 
the trapezius, to the transverse process of the first cervical 
vertebrae. 

A etemo-hyoideus profundus (Fiirbringer) passes from the 
anterior edge of the sternum to the hyoid. It is not distinctly 
continuous with any of the preceding. It is interrupted by 
the clavicle ; and the hinder part of it is called episterno-clavi- 
cularis by FUrbringer. A few fibres passing from the sternum 
to the coracoid constitute a aterno-coracoideua, 

SternO'Costalia [triangularis sterni) radiates, from a tendinous 
attachment to the hinder surface of the sternum, upon the 
deeper surface of several of the foremost ribs : it is in the 
stratum of the transversalis. There is no trace of diaphragm. 

Scaleni are continuations forwards of the muscles between 
and upon the ribs to the cei-vical transverse processes. One slip 
reaches the lateral part of the occipital. 

Latissimus dorsi {trapeziums of Fiirbringer) is small and 
thin, arising from the spines of five or six vertebrae behind the 
girdle. It is in the same plane with the trapezius but does 
not reach it. Its fibres converge ; and the flat muscle fonned 
by them passes over the dorsal . surface of the scapula, lying 
upon the scapular cartilage but deriving no fibres from it, and 
is inserted into the hinder edge of the clavicle. 

Near the scapula it is joined by a bundle from the rectus 
abdominis, which is a serial continuation of the bundles passing, 
from the lateral part of the rectus, to the ribs near the vertebrae. 

^ Nob. 2 and 8 are described by Biidinger as Pectoralis minor, by Fiirbringer 
ftB itertio-costo scapularii. 



•i 



I 



ON THE DISPOSITION OF MUSCLES IN VERTE- 
BRATE ANIMALS. By Peofessor Humphry*. 

The locoraotory system of a vertebrate animal consists, funda- 
mentally, of a successional series of alternating transverse 
skeletal and muscular planes which extend nearly through the 
outer wall of the animal. I say nearly through, for in and 
around the axial line, and in the immediate mesial plane, 
this alternation does not exist. The skeletal elements here 
form a continuous structure composed, in the axial line, 
during the early foetal state, and persistently in the Lancelet, 
of a simple cartilaginous notochordal streak, but usually, at a 
later period, of cartilaginous or osseous pieces articulated directly 
together or connected by ligamentous material. This vertebral 
column extends throughout the whole length of the animal; and 
cartilaginous or osseous processes run out from it, or in con- 
nection with it, more or less transversely, and serve as levers 
to aid the muscles besides fulfilling other purposes. It is 
obvious, however, that these processes must, in proportion to 
their length and unyielding character, limit the range of 
movement of the axial pieces upon one another and so lessen 
the flexibility of the animal. Accordingly, in the circumfer- 
ential regions, especially in the directions in which move- 
ment is most required, the skeletal parts are not osseous 
or even cartilaginous, but are membranous and composed of 
fibrous plates extending from the axial osseous or cartilaginous 
structures to the skin. This may be the case throughout the 
whole of the muscular stratum as in the Lancelet, where the 
membranous septa extend from the skin down to the noto- 
chordal sheath and blend with it. The muscular planes occupy 
the intervals between the osseous processes and between the 
fibrous plates, the latter being continuous with the osseous 
axial pieces and their processes. 

^ 1 The anatomical points mentioned in this paper, miless oiherwiBe specified, 

r are aU from notes, published in the Joum. of AnaL, or in manuscript, of dissec- 

tions made by Mr Anningson or myself. I have not been able to investigate the 
now copious literature of the subject so much as I could have wished. 



294 PROFESSOR HUMPHRY. 

This arrangement is found most distinct in the simplest 
parts of the lower vertebrates, as throughout the Lancelet 
and in the caudal region of Fishes and Urodelans. The struc- 
ture of that region, with its longitudinal divisions caused by 
the dorsal, neural and lateral septa, and its transverse divisions 
caused by the transverse septa, has been described in the 
Cryptobranch (p. 3)\ 

The transverse skeletal planes, membranous, cartilaginous 
and osseous, are sometimes called ' sclerotomes ' and the trans- 

^ I Bhonld observe that the angular spaces left between the longitudinal divi- 
sions are not always, as in Cryptobranch, occupied by fat. In the Fish the 
dorsal and ventral furrows are often partly occupied by the dorsal and anal fins 
(see pp. 256, 277) ; and the lateral furrows are commonly occupied by muscular 
fibres which bear the transverse septa, but which are more closely connected with 
the skin, and peel off with it more easily than the rest of the lateral muscle. 
These fibres are more vascular than ordinary muscular fibres ; and in a piece 
which I examined from a Dace they contained more oil than the other mus- 
cles. Stannius {Handbuch der Zootomie, ii. 112) says that they, in addition, 
present microscopically the appearance of tissue in process of conversion into 
muscle. I did not find that to be the case. With the exception of the excess 
of oil, they presented the usual microscopical characters of striped muscle. 

Stannius (ii. 93) uses the terms 'epaxonisch' and 'hypaxonisch' to indicate 
the muscles above and below the axial vertebral line. Huxley {The Anatomy of 
Vertebrate AnimaU) uses the terms * episkeletal' and * hyposkeletal' to indicate the 
muscles situated respectively above and below the endoskeleton, and developed 
from above and beneath the protovertebras. Strictly speaking, however, all the 
muscles are interskeletal; forasmuch as the intermuscular septa exteM, from 
the transversalis fascia to the skin, through the entire thickness of the muscular 
layer which appears to be primarily and essentially one, and which is, for aught 
we know, all developed from the same embryonic protovertebral stratum ; and 
the skeletal tissues undergo chondrification and ossification in certain parts only 
and in certain planes. The parts in which these changes take place are chiefly 
in and near the axial line. The plane in which chondrification and ossification 
occur is almost exclusively the middle one ; and ossification without chondrifi- 
cation — i. e. from membrane — is most frequent in the superficial muscular 
plane ; though it may take place much deeper, as in the instances of the para- 
sphenoid, or sub-basal, bone and the vomer. I am not here speaking of the epi- 
dermal, or superficial dermal, tissues which may also be changed into homy 
matter, cartilage or bone. The ossifications in this epidermal layer are as dis- 
tinct from those in the outer, or subcutaneous, muscular plane as these are from 
the chondrifications and ossifications in the middle muscular plane, or more so ; 
and I do not think this distinction has been quite sufficiently kept in mind. It 
must not be forgotten that the ' cartilaginous,' the * membranous,' and the * epi- 
dermal' ossifications may be blended, and that the two former are particularly 
liable to be so. Indeed, those that begin in cartilage are usually enlarged by the 
addition of membrane bone. Histologically, there is perhaps no essential 
difference between the two. 

It is, I think, far better to reserve the prefix * epi ' to designate the bones — 
membrane bones — which are formed, usually, thou^ as above mentioned not 
always, in the superficial muscular or subcutaneous strata, and thereby distin- 
guish them from the subjacent cartilage bones with which they are often closely 
related. Thus the 'epistemals' and the 'epicostals' are indicated to be bones 
found in the tissue overlying the sternum and the costsB ; and the * epiooracoids * 
are bones or bony plates formed over, or upon, perhaps blended with, the 
ooraooids. 



V 



i 
i 



MUSCLES IN VERTEBRATE ANIMALS. 295 

Terse muscular planes between them 'myotomes/ Very rarely, 
however, are the planes truly transverse. They commonly 
slant with more or less obliquity, backwards or forwards, 
from the axial line towards the circumference. An additional 
and more perplexing element of confusion is imported by 
the membranous or fibrous portions of the sclerotomes — the 
intermuscular septa as they are called — ^not preserving a uni- 
foim direction, but slanting first one way then another, first 
backwards and then forwards, or vice versk, as they are 
traced from the upper or dorsal edge of the animal. Thus 
they acquire a more or less waving or zigzag line ; and their 
supei-ficial margins come to deviate considerably from the 
lines of their deeper margins and the lines of the osseous 
vertebral processes to which they are attached. Moreover 
the several curves or angles so formed may be produced to 
a great length. Especially is this the case near the mesial 
line, above — ^in the *mesio-dorsal' part of the lateral muscle. 
Here the angles of the septa are often prolonged to a con- 
siderable distance; and in some Fishes (Bream and Dog-fish) 
they are, near the surface, thickened into tendons with the 
muscular fibres on the two sides of each passing obliquely 
between it and the adjacent septa, which has the effect of 
allowing the traction of a large number of muscular fibres 
to be brought to bear upon a given point or points (p. 271, 
Fig. 28). Where this occurs the intermuscular septa cease 
to run directly into the skin or to retain their close con- 
nection with it; for the cutaneous terminal filaments become 
converted into loose areolar tissue. Hence the tendinous septa 
as well as the intervening muscular planes, near the dorsal 
median line, in the Fishes above mentioned and in others, 
are allowed to glide with greater freedom beneath the skin 
than are the more lateral and ventral muscles, which are 
bound to the skin by the intermuscular septa running from 
the axial osseous structures into it. 

A further change consists in the isolation of the pro- 
longed and tendinous superficial parts of the septa from one 
another, and, to a greater or less extent, from their muscular 
contingents, as well as from the deeper and the superficial 
structures; so that they run alone to their destinations and 



296 PROFESSOR HUMPHRY. 

admit of traction without hindrance. This is effected by the 
conversion of the surrounding connecting structures into loose 
tissue, as just mentioned in the case of their isolation from 
the skin. Such a condition we find developed to perfection 
in the numerous delicate muscles and tendons which lie 
along the sides of the dorsal spinous processes of Snakes. 

Thus, by comparatively simple stages of transition, the 
elongated longitudinal dorsal muscles are brought into rela- 
tion with, or reduced from, the simple primary transverse 
muscular strata; and it is to be remarked that while this 
change takes place, usually in some degree and in many 
instances to the extent I have mentioned, in the upper or 
* mesio-dorsal' part of the lateral muscle, the transverse dis- 
position is commonly maintained through the visceral region, 
at any rate in the opposite and lowest, or 'mesio-ventral* 
part of the same muscle. Witness the transverse direction 
of the septa when they are persistent in the recttia ahdominis 
and in its extensions into the neck and the tail. 

So much for the variations in the fibrous plates or septa 
of the skeletal structure. Then, with regard to the muscles be- 
tween them. These, in the simple condition, occupy the in- 
tervals between the septa and are composed throughout of 
fibres passing, antero-posteriorly, from one septum to another. 
Through the medium of the septa the fibres of one compart- 
ment are connected with those of the adjacent compartment, 
and also with the subcutaneous fascia and the skin, as well 
as with the osseous structures of the skeleton and the inter- 
nal or transversalis fascia. I have already mentioned that 
where the septa are very oblique the muscular fibres between 
them usually take an oblique direction also. 

Sometimes the fibres at one part of the thickness of the 
lateral muscle take an oblique direction different from those 
at another depth. This alone is sufficient to cause a cleavage 
into planes; and the cleavage usually extends through the 
septa as well as between the muscular fibres, so causing the 
separation of one or more superimposed muscular sheets which 
are thus rendered capable of moving upon one another \ 

^ The cleayage of a muscle into two planes, in oonsequenoe of a difference in 
the direction of its snperfioial and deep fibres, is well illustrated in the pectoralU 



I 
I 
t 



MUSCLES IN VERTEBRATE ANIMALS. 297 

The Cryptobranch (p. 10) and the Lepidosiren (p. 256) offer ex- 
amples of the cleavage bein^ confined to the muscular elements 
without involving the septa. When the cleavage involves the 
septa these may remain (p. 258). Commonly they disappear, 
more or less completely, throughout the thickness of the muscu- 
lar substance. A continuity, or ankylosis of the muscular fibres 
of the several compartments is thus established ; and all trace 
of the primary, transversely segmented, myotomic arrangement 
is obliterated*. 

Even without an alteration in the direction of the muscu- 
lar fibres a cleavage into superficial and deeper planes may 
take place; and it may take place completely or partially. 
Thus a superficial stratum of muscular fibres, having the same 
or a different direction from those beneath them, may be 
quite severed from the deeper strata. It may retain here 
and there connection with the skin only, so forming, as is 
the case with portions of the pannicle, a purely 'cutaneous' 
or as commonly designated 'subcutaneous' muscle ; or it may 
retain, at one or more places, a connection with the deeper 
strata of the muscle or, through remnants of the transverse 
septa, with the osseous skeleton. Thus it may be, as nume- 
rous varieties of the pannicle shew, a 'musculo-cutaneous' or 
'osseo-cutaneous' muscle*. Lastly, a given layer may be severed 
from the deeper strata in a part only of its length, and may 
retain a connection, through the septa, which as already 
mentioned are equivalent to tendons, at both ends, either 
with the subjacent muscle or with the subjacent skeleton; wit- 
ness some of the dorsal muscles, the muscles passing to the 
girdles, and many others. That is to "say, a superficial or cuta- 
neous stratum may be segmented more or less completely 
firom the rest of the muscular system; and various strata of 

major of Cyolothnrns {Joum, AnaL iv. 25), and still better in that of the Wild 
Cat, in which there are no less than four layers. 

^ This is exemplified even in the Myxinoid fish Bdellostoma; and the con- 
trast-between the absence of inscriptions, or 'ligamenta intermtucxdaria,* in the 
oblique muscles, and their presence in the straight muscles, is well shewn in 
Tab. I. accompanying Miiller's well-known paper, Abhandl. Berlin. Akad, 1884. 

* In Pteropus (Joum. Anat. ui. 299), the cutaneous muscles are connected with 
the skull, the sternum, the coracoid, the pelvis, and the femur, thus presenting 
unusually numerous and good examples of 'osseo-outaneous' muscles. They 
are also remarkably well developed and have several osseous connections in the 
Hedgehog. 



298 PROFESSOR HtJMPHRT. 

the remaining deeper portion may be more or less completely 
segmented from each other. • 

It is a very common thing for the muscular fibres to miBS, that 
is, to pass over or under, one or more septa, as in the case of the 
erector spine and fche subcostals. In these instances, and others of 
the like kind, the muscular fibres are continued through, or are 
ankylosed through, the septal tissue which has disappeared as such, 
owing to the embryonic tissue of the intermuscular septa undergoing 
the same histological change as the muscular parts. See p. 301. 

A muscular plane, or any part of it, may also be divided 
longitudinally into portions or sectors by cleavages similar to 
that by which itself was separated from the strata above or 
beneath it. 

Thus from the simple primary, transversely segmented, late- 
ral muscle, on either side of the animal, the various muscular 
forms may be elicited. Moreover the diflferences in the muscu- 
lar systems of different animals, and in different though serially 
homologous parts of the same animal, may in great measure be 
explained by variations in the number of the strata or of the 
sectors, or by variations in the depths at which the several 
strata have been detached, or in the points at which the several 
sectors have been separated. Hence, although general coito- 
spondence may be indicated, precise homology must not be too 
closely pressed. 

The processes which I have mentioned miay be recapitulated 
as follows : 1, varieties in the inclination and direction of the 
septa; 2, prolongation of the angles of the septa caused by 
increased inclination and flexure ; 3, separation of the thus 
prolonged septa with their appended muscular fibres, indivi- 
dually or in groups, into independent muscles ; 4, variation in 
the direction of the muscular fibres; 5, cleavage into planes 
and into sectoi*s reaching to various depths ; 6, fusion or anky- 
losis of the muscular segments by the establishment of conti- 
nuity through the septa. 

In the three highest ordei-s of vertebrates, if we exclude the 
vertebral processes, the ribs, the hyoid and the limb girdles, which 
are ossifications in the deeper parts of the septa^ and the tendons 
of the dorsal muscles^ which are modifications of the 'septa, the 
intermuscular septa are represented only, or chiefly, by the 



<•■ 



MUSCLES IK YERTEBBATE ANIMALS. 299 

inscriptions in the rectus abdominis, biventer cervicis, digastric 
and omo-hyoidy by occasional inscriptions in the stemo-hyoid 
and stemO'thyroid, by the clavicle or the inscription which, 
in carnivora and some others, is substituted for it between the 
trapezius and the deltoid, and by Poupart's ligament. 

The fibres in these animals retain their simple antero- 
posterior direction between the successive skeletal septa in the 
instances of the interspiTiales and intertransversales and in the 
recti-abdominis, though in the last some of the septa have been 
obliterated. In the semispinales the fibres run obliquely be- 
tween the successive skeletal structures and also in the inter- 
costales. In these last, the direction of the obliquity varying at 
two parts of the depth between the successive ribs, two strata 
are formed resembling the two strata of external and internal 
oblique between the successive ribs and septa in the abdomen 
of Cryptobranch (p. 10). These and other variations will, how- 
ever, be mentioned as we proceed. 

Reverting to the plan of the construction of the locomotory 
system of a vertebrate animal, we have found it to consist of a 
series of transverse, alternating, skeletal and muscular planes or 
discs, traversed by an antero-posterior axial line, the whole 
being enclosed in an external fascial sheet which is also part of 
the skeletal system and which lies beneath the skin. A trans- 
verse section shews that each disc is in two lateral halves 
applied, as it were, upon the axial line which is formed by the 
notochord or the vertebral bodies. Each half constitutes one 
side of the animal Owing to the obliquity with which the 
planes or discs slant from the axial line, a transverse section 
passes through both the muscular and the skeletal planes and 
shews their relative disposition. Such a section indicates the 
locomotory system of an animal to consist on each side of a 
fascial, or skeletal, tube enclosing a muscular cylinder, which 
last is traversed by transverse or oblique, fascial or skeletal 
planes. The approximated sides of the skeletal tubes are 
flattened against each other, above and below the axial 
line; and becoming fused here in great part of their extent 
they form mesial septa passing, dorsally and ventrally, from 
the axial line. 

This is well illustrated by the section of the tail of a Tad- 



300 PROFESSOR HUMPHRY. 

pole (Fig. 41) *, in which the fascial walls on either side, ascend- 
ing and descending from the notochord and enclosing the neural 
and haemal canals, meet and so form the dorsal and haemal 
septa. These run peripherally into the upper and lower mem- 
branous edges of the tail, where they are joined by the fascial 
layers from the exterior of the lateral muscle. Thus each mem- 
branous edge of the tail is composed, essentially, of four sheets 
— the two mesial sheets which are blended together and form 
the septum, neural or haemal, and the two external, or lateral 
sheets which are derived from the superficial covering, or fascia, 
of the lateral muscle. It is interesting to observe that at the 

^ The specimens of Fseadis Tadpole, from which the drawings were made, 
were kindly sent me by Mr MiTart. 

Description of Figures of the Pseudis Tadpole (PL XV.). 

Fig. 89. — Side Tiew of the animal, shewing the lateral muscle of the tail. This 
at the hinder part is in interrupted fragments, which are serially arranged in 
rows with wide septa between the rows. Further forwards the muscular fibres 
are more developed, filling in the rows and diminishing the intervals between 
tiiem. A pin (C) is placed in the cloacal opening ; another, above B^ is in the 
branchial opening. Above, and in front of the branchial opening, is the eye. 
The small hind limb is seen on the hinder wall of the visceral cavity, or rather 
in the furrow between it and the tail, above L, 

Fig. 40. — The visceral cavity laid open in a specimen more advanced where 
the cavity is more elongated. The transversely marked ventral portions of the 
lateral muscle are seen extending, from the tail, along the sides of the bodies of 
the vertebrsB, and projecting into the visceral cavity, with an interval between 
them leading down to the haemal passsige which has ceased to be a covered 
oanal, and is merely a channel. The waU of the visceral cavity has been 
turned back ; and the commencing development of its muscle is shewn, in iso- 
lated tracts, by fine muscular fibres arranged in rows between broad, white (sep- 
tal) lines. 

Fig. 41. — A transverse section at about the middle of the tail. The large 
central circle is the notochord, with a small neural canal above and a small 
hasmal canal below. — J). P,^ the dorsal, or neural, membranous fringe. — V. F,, 
the ventral, or htemal, membranous fringe. The neural septum extends from 
the notochord up the middle of the one, and the hiemal septum down the mid- 
dle of the other. The sides of both are formed by extensions of the membrane 
covering the lateral muscle. Tbe base of each, between the mesial septum and 
the external membrane, is occupied by soft succulent tissue which is separated 
from the lateral muscle by an offset from the external enveloping membrane. 

Fig. 42. — A section through the animal, made at the back part of the vis- 
ceral cavity, and seen from in front. D. F., the dorsal fringe with the neural 
septum descending, through its middle, to the neural canal and the notochord. 
Beneath the latter is the naemal passage at the bottom of a deep channel be- 
tween the lateral muscles of the two sides, or, rather, between the plates of the 
haemal septum, which have not coalesced, and which separate the hsmal chan- 
nel from the lateral muscles. Beneath this channel is, (7, the cloacal tube. A 
is the wide posterior cul-de-sac of the abdominal wall, with the muscular fibres 
beginning to be developed in rows between the paler lines, which indicate the 
future septa. 

Fig. 43. — A section farther forwards. A^ the abdominal, or visceral, wall 
ascends, is reflected, higher on the sides of the lateral muscle, reaching to the 
dorsal part of that muscle. The hsamal channel is wider and forms part of the 
visceral cavity. 



MUSCLES IN VERTEBRATE ANIMALS. 301 

extremity of the tail (Fig. 39), as well as along its upper and 
lower edges, the membranous sheets alone exist, the muscular 
fibres being absent ; so that the mesial and external membra- 
nous plates are in contact. A little in front of the extremity of 
the tail the muscular fibres begin to appear, in patches, separa- 
ting the external from the mesial plates. The patches are in 
broken rows ; still they succeed one another in serial order. Gra- 
dually, as they are traced forwards, the rows are filled up and the 
membranous interspaces between them become reduced to the 
narrow intermuscular septa; while the muscles, increasing in 
thickness, acquire the semicircular or ovoid form which is seen 
on either side of the median line in a transverse section (Fig. 
41). It is thus perceived that the membranous, or intermus- 
cular, element is the first formed, that the muscular fibres are 
produced, or added in it, that the intermuscular septa and 
fascial sheets are remnants of it, and that these give way, in 
greater or less degree, before the force of muscular development. 

Usually, in other animals, ossification takes place, to a 
greater or less extent, around the notochord, giving rise to the 
bodies of the vertebrae, and extends into the ascending and 
descending mesial laminae, forming the neural and haemal 
arches and spines. In Fishes (Figs. 31, 32 and 33) it also often 
extends into these laminae, where they stretch, like the mem- 
branous fringe of the Tadpole's tail, beyond the confines of the 
lateral muscla Thus are formed the fin-ray bones, which, like 
the septa, are double, actually or potentially; and muscular 
fibres are formed upon them. These have the same relations 
to the membranous laminae, and the same segmentation, as the 
fibres of the lateral muscles, and are, indeed, extensions of 
them, though they take a different direction, and are, con- 
sequently, segmented from them (p. 257). 

In front of the tail, in the visceral region, the mesial 
haemal laminae are kept apart; and the visceral cavity is 
formed between them. In other words, they are spread out 
over it, and form the fascia transversalis. This separation of 
them may, in the hindmost region of the visceral area, be con- 
fined to the marginal part of the haemal septum: thus, in 
Fig. 39 of the Tadpole, the cloacal tube and aperture are seen to 
lie at the junction of the membranous caudal fringe with the 

VOL. VL 21 



302 PBOFESSOR HUMPHRY. 

abdomen ; and they are the result of a want of adhesion of the 
mesial laminae which form that fringe. In the Fish the cloacal 
fissure, or separation, of the mesial laminae, usually extends 
deeper into the region of the haemal spines, which are, accord- 
ingly* bifurcate and arch transversely over this part of the 
visceral cavity instead of, as in the tail, occupying a median '^ 
position, and each pair being fused into a single process. In 
front of the cloacal region, and through the rest of the visceral 
cavity, with occasional exceptions near the head, the separation 
of the mesial laminae extends dowu to the vertebrae, splitting 
the haemal arches of the two sides quite asunder, and laying the 
haemal canal open to the visceral cavity. Towards the fore part 
of the visceral cavity the mesial laminae and the haemal arches 
are pressed so far upwards upon the vertebral bodies that the 
arches come to occupy the place of ribs. 

The wall of the visceral cavity is thus formed by the sepa- 
rated and expanded haemal plates, and not merely by those 
parts of the plates upon which, in the tail, the lateral muscle 
is formed, but by those parts also which form the membranous 
haemal fringe in the Tadpole's tail, and in which the subcaudal ^ 

or anal fin is formed in the Fish. In the Tadpole it is chiefly '"'^ 

this latter, membranous, fringe-like part which becomes ex- 
panded ; for the lateral muscle (the ventral portion of it) is 
continued, without much alteration, from the tail, forwards, 
through the abdomen \ as represented in Figs. 4-0, 42 and 43. 
These figures further shew the manner in which the membra- 
nous part is spread out and is reflected upon the sides of the 
lateral muscles, so as to give greater space for the contents of 
the visceral cavity; and they shew the mode in which the 
development of the abdominal muscles is commencing in the 
thickness of the wall by broken serial rows of muscular fibres 
with intervening septal lines, upon which the muscular trans- 
formation gradually encroaches, in the same manner as the 
development of the lateral muscle is progressing at the end 
of the tail (Fig. 39). Further, in Fig. 40, an extension from 
the sides of the lateral muscle is seen to be proceeding into the 
hinder part of the abdominal wall. In the Fish, the part of , ^ 

^ It is reduced and becomes the qvadratui lumborum in the Frog. 



MUSCLES IN y£RTEBRAT£ ANIMALS. 303 

the hsemal plate in which, in the tail, the lateral muscle is 
developed becomes, in the abdomen, more expanded than it 
does in the Tadpole, and forms, at any rate, the ventrolateral 
(F. L., Fig. 31) portion of the body-wall. It is not improbable 
that the part in which, in the tail, the anal sub-caudal fin is 
developed is continued forwards into the abdomen as the mesio- 
ventral {M, V., Fig. 31) portion of the body-wall. Of this, how- 
ever, I cannot be sure, the two parts ( F. L. and M. L.) being, 
as we might expect from their development in one continuous 
hsemal plate, blended into one 'lateral' muscle \ There can, 
however, be little doubt that, as pointed out by me, in a paper 
" on the Homology of the Mesial and Lateral Fins of Fishes,'' in 
this Journal, Vol. V., the ventral and pectoral fins and their 
muscles are formed from the same serial elements as the sub- 
caudal or anal fins and their muscles. 
I will now consider, briefly, 

The Dorsal Muscles of the Trunk, 

that is, the muscles situated above the lateral septum. These, 
though numerous, in accordance with the number of the 
vertebrae that require to be moved upon one another, do not 
present much variety or much interest in diflferent animals. 
In the Fish the dorsal mass from which they are derived indi- 
cates a division into an upper or 'mesio-dorsal' and a lower or 
' latero-dorsal' part. This division corresponds on the whole 
with that into the spinalis and hngxssimus dorsi and the sacro- 
Iwmhalis parts in higher animals. Such incipient longitudinal 
segmentation is less marked in Cryptobranch, Perennibranch, 
and Lepidosiren, than it usually is in Fishes. It is seen again 
in the Salamanders. 

The dorsal muscles may be arranged, as follows, in two 
divisions : 

First, those in which the fibres retain the primary antero- 
posterior direction and pass between corresponding parts of con- 
tiguous or distant vertebrse. Those connected with contiguous 

^ Snpposizig this yiew to be oonreot, the ttenuJ rilvi and the Btenmm would 
be serial representatiyes, not of the hsmal processes bnt, of the osseous ele- 
ments of the snboandal fln— the fin ray-bones — of the Fish. 

21—2 



304 PROFESSOR HUMPHRT. 

vertebras are commonly designated in accordance with the parts 
of the vertebrsB between which they pasa Thus they are called 
* Interapmales'y ' IrUertransversalea't and * Interobliqui' or 'In- 
teracces8or%i\ The foremost of them is the Rectus capitis pos- 
ticus minor. They are developed in proportion to the mobility 
of the parts between which they pass, that is, they are most ^ 

developed in the neck and loins, and least in the back. In the 
back the intertransversales and interspiTudes are sometimes 
merely ligamentous. In the more superficial members of this 
series the fibres sometimes leap over one or more segments to a 
distant point. This is caused by the superficial fusion of two 
or more septa, owing to the non-development of muscular fibres 
between them, or by the superficial obliteration of one or more 
septa from the extension of muscle*development through them, 
producing, in the one case, an elongation of septum or tendon, 
and, in the other, an elongation of muscle. In this manner are 
formed the spinalis dorsi and spinalis colli and the several parts 
of the erectores spines. The last, it may be observed, often 
overlap, to some extent, and are connected with the veAtral 
parts of the skeleton, viz., the iliac bones and the ribs, parts, 
that is, which lie beneath the lateral line. "^ 

The second division includes the muscles in which the fibres 
have an oblique direction and pass between non-corresponding 
parts of contiguous or distant vertebrae, for instance, from 
transverse process to spine, or from spinous process to trans- 
verse process. The obliquity is therefore in two directions, 
giving rise to two sets, the fibres of which cross one another. 
In the one set the fibres pass from spinous processes outwards 
and forwards to the transverse processes; and in the other set, 
which is on the whole in a deeper plane, the fibres pass from 
the transverse processes inwards and forwards to the spinous 
processes. The more superficial, spino-transverse or ot^^wardly 
directed, set comprises the obliquus capitis inferior, and the 
splenius capitis and coUi, . The deeper, transverso-spinous or 
tnwardly directed, set comprises the obliquus capitis superior, 
the complexus with the biventer, the semispinalis colli and dorsi, 
the mvitifidiLS and rotaiores spines. .^ 

The dissections of Cryptobranch (Fig. 20) and of Lepidosiren 
(Fig. 25) show conclusively that the temporal muscle is a pro- 



irUSCLES IN VERTEBRATE ANIMALS. 305 

longation of the dorsal muscle forwards from the neck, beside 
the cranial neural arches, over the suspensorium, to the lower 
jaw, and that the temporal fascia is the continuation of the 
fascial aponeurosis of the dorsal muscle. Thej render it pro- 
bable that the masseter and external pterygoid, and also the 
ocular muscles, are an extension of the same series. According 
to this view the temporal ridge which, in most vertebrates, 
shuts off the masticatory muscles from the rest of the dorsal 
system, may be regarded as an ossification of, or an ossification 
extending into, a transverse intermuscular septum, an ossifica- 
tion, that is, in the superficial stratum of the dorsal muscle 
passing upon the head. 

The ligamentum nuchce and the interspinous ligaments, as 
well as the ligamentous bands tying the skin to the several 
spinous processes, are modifications of the longitudinal median 
septum which I have already (pp. 254, 300) described. 

The modifications of the dorsal muscle in the tail do not 
require any special description. It may be sufficient to remark 
that in cases where the pelvis is absent or rudimentary, as in 
the Porpoise, the portion of it called erector spince is continued 
uninterruptedly from the lumbar to the caudal region. 

The Ventral Muscle 

is subject to much greater modifications in the different regions 
of its course than is the dorsal. In the hinder part of the tail 
it much resembles the dorsal muscle of the same part; but, an- 
teriorly, the symmetry between the muscles above and below 
the lateral line is destroyed by the expansion of the ventral 
muscle over the visceral cavity, by the formation of the limb- 
girdles in its substance, and by its relation to the limbs. Tra- 
velling forwards it first comes into relation with the openings 
of the alimentary, urinary, and genital organs, and detaches 
muscles to them. It then encounters the pelvis and hind limb, 
which, more or less, interrupt and make demands upon it. 
Next it is expanded, and the direction of its fibres is modified, 
by the visceral cavity. Then the shoulder-girdle and fore- 
limb, the branchial and hyoidean apparatus, the larynx and 
pharynx, the lower jaw and the face necessitate modifications 



306 FROFESSOB HUMPHBY. 

to meet their several requirements, which vary in different ani- 
mals, and which lead to almost infinite divemties in the dispo- 
sition of the several parts of the muscle. 

The ventral muscle, and this is important^ not unfrequently over- 
laps the dorsal muscle. Such is described by Miiller to be tlie case in 
Bdellostoma^ j and superficial fibres or strata of it are often pro- 
longed into the tissue over the doi'sal muscle, so reaching the dorsal 
spines, or are continued as distinct muscles to this region (see pp. 13, 
260, and Fig. 24). This occurs especially towards the fore part of the 
trunk, and is exemplified particularly in Snakes (Fig. 44) ; and this 
extension contribute to the formation of the superficial muscles of the 
neck, throat and face, as well as to those of the limb-girdle and limb. 

While the superficial strata of the ventral muscle thus overlap the 
dorsal muscle, and reach to the dorsal median line, the deeper strata, 
as represented by the intercostals and the ribs, are commonly, to 
some extent, overlapped by it; so that the lateral margin of the 
dorsal muscle is received between the layers of the ventral muscla 

In the simplest condition, as in the ordinary teleostean fish, 
the ventral muscle does not undergo much change in its dif- 
ferent parts. The two fascial haemal plates, as has just been 
shewn, which line the apposed sides of the ventral muscle, in 
some parts of the tail coalesce beneath the haemal spines, and 
form a median osseo-membranous septum, extending from 
the haemal spines to the skin. At other parts they are sepa- 
rated by the caudal or anal fins. At the foremost part of the 
tail they are also separated; and the commencement of the 
visceral space exists between them, but walled off from the 
haemal space by the haemal arches still bridging over the latter. 
Further forward the separation is greater, the osseous haemal 
arches disappear, the haemal and visceral cavities are laid into 
one, or are divided only by membrane. The ventral muscles 
of the two sides are pushed asunder; and the separated mem- 
branous haemal plates which line them are named the fascia 
transuersalis. Ossifications in the inner parts of the transverse 
septa which pass from the fascia transversalis, through the lfi.te- 
ral muscle, to the external fascia and the skin, form the ribs. 

^ AhhandL der A had. Berlin, 1834, b. 245. It may be observed also that 
in Bdellostoma (Lc. p. 246) the saperfioial fibres of the ventral musde, in 
the interval between the head and the hinder opening of the giUs, cross the 
median line and are oontlnued from either side upon the surface of the ventral 
muscle of the opposite side, where they are lost in the superficial fascia. This, 
however, is qnite exceptional. 



V 



MUSCLES IK VERTEBRATE ANIMALS. .307 

The latter thus correspond serially, or nearly so, to the haemal 
spines, split and pressed asunder, and to the sides of the haemal 
arches. If an interval is left between the ribs and the trans- 
versalis fascia, in which muscle is developed, it constitutes the 
transversalis muscle, or it may constitute the levatores, or, more 
commonly, the depressor es costarum^ according to its position*; 
and when this muscular sheet extends beneath the bodies of 
the vertebrae it constitutes the svbvertebral rectus (p. 11). 

The mesial edges of the ventral muscles, inferiorly, are se- 
parated and covered by fascia ti^wnsversalts continued round 
each into the external fascia. Behind, there is an interval be- 
tween the contiguous edges of the ventral muscle, thus covered 
by fascia, which permits the passage of the alimentary tube to 
the anus. A layer of the fascia (Fig. 23, G) accompanies the 
tube and binds it to the skin; and some muscular fibres may 
be developed around it forming a sphincter. 

In front of the anus the pelvic bones lie between the fascia 
transversalis and the ventral muscle, or in the substance of the 
deeper layers of the ventral muscle, in the same plane, that is, 
as the ribs. The fins project between the mesial edges of the 
ventral muscle; but marginal portions, or (Bream) a larger 
amount of the deep stratum, of that muscle are attached to the 
pelvic bones, constituting a *retra,ctor' or * protractor* ; and 
more superficial portions constitute what are sometimes called 
'carinales' muscles. In some Fishes (Ceratodus) portions of 
the ventral muscle extend upon both surfaces of the fin. The 
pectoral fin projects at a more lateral point; and the ventral 
muscle is attached more largely to its girdle which, like the 
pelvic bones, occupies the same relative position, with reference 
to the thickness of the abdominal wall, as the ribs. From it 
the ventral muscle is continued forwards to the hyoid and the 
head. 

1 In the Bream, Dace, and some other Fish, there is a sheet of mascnlar 
fibres passing from the yertebrie forwards and downwards internal to the ribs, 
that is, taking a direction corresponding to the internal intercostals and sub- 
costals of mammals, and to the internal intercostals, depressores eottarum and 
transvenalis of Cryptobranch. In Ceratodus (p. 280) I haye described tendinous 
fibres taking the same direction. Some are attached to the ribs nearest to their 
point of origin, others to more distant ribs. The nerves are external to this 
sheet. Towards the fore part of the body of the Dace fibrous bands take the 
place of these suboostals. 



30S PROFESSOR HUMPHRY. 

The median third or half of the ventral muscle is not tin- 
frequently distinguished from the remainder in Fishes by the 
mere transverse direction of its septa. In some, as the Dog- 
fish (Fig. 28), it is segmented from the remainder, in great part 
of its extent, and overlaps it. In Mammals also it is separate, 
as rectuSf and is enclosed in a sheath formed by prolongations 
of the lateral portions of the ventral muscle, or their tendons, 
to the mesial line. Its continuity with the lateral parts of the 
ventral muscle has been shewn in Cryptobranch (p. 11), and in 
Lepidosiren (p. 256). In Cryptobranch, however, it must be 
observed that while the more superficial or oblique strata of the 
ventral muscle are continued into the rectuSy the deepest or 
transversaiis stratum, separating itself from the others, passes 
upon the deeper surface of the recttis to the middle line ; and 
in Mammals the rectus is ensheathed by tendinous extensions 
from the deep, or peritoneal, and the superficial, or cutaneous, 
divisions of the ventral muscle which pass, upon its peritoneal 
and cutaneous surfaces, to the middle line. Though we are in 
the habit of thus referring the walls of the sheath of the rectus 
to the expansions of the lateral parts of the ventrcd muscle, we 
may, with equal right, assume them to be deep and super- 
ficial strata of the median part of the ventral muscle, that is, 
derivations from the rectus itself. And as, on the one hand, 
they retain their continuity with the lateral parts of the muscle, 
so, on the other hand, do they retain their connections with the 
rectus, or middle stratum of the same, through the transverse 
inscriptions which extend from it into them. 

As already intimated the ventral muscle is disposed in three 
chief planes or strata which are in variable degrees distinguishable 
in different animals and in different parts of the same animal. Of 
these, the middle — or internal oblique — ^plane in that in the septa of 
which the pieces of the true or cartilage skeleton are for the most part 
developed : the external— or external oblique — stratum is conneeted 
with the external fascia and the skin, and is that in which the pieces 
of the dermal or membrane skeleton are for the most part developed : 
and the internal~-or transversaiis — plane is connected with the inter- 
nal or transversaiis fascia and the viscera^ much in the same way as 
the external plane is connected with the external fascia and the skin. 
The connection with the viscera, however, is in great part interrupted 
by the formation of the visceral cavity. Piimarily all the planes are 
continuous or, rather, formed one plane from the skin to the visceral 



MUSCLES IN VERTEBRATE ANIMALS. 309 

tube. It remains so in the Leech. In the Earthworm the transverse 
septa still connect the visceral tube and its muscles with the dermal 
sheet and its muscles. In higher animals the visceral cavity inter- 
rupts the septa ; and they are restricted to the dermo-muscular layer, 
except near the oral and anal apertures, and in the situation of the 
diaphi-agm or diaphragms, where the inner layer or layers of the 
dermo-muscular sheet bend in towards and come into contact with 
the visceral sheet of the animal. In the Fish the dermo-muscidar 
sheet is, for the most part, in one undivided plane, with the septa 
extending throughout it and connecting the skin with the fascia 
transversalis. In Batrachians the muscular part is more separate 
from the skin on the one side, and the fascia transversalis on the 
other; and the stratification of the muscular part into the three planes 
above-mentioned is commencing. 

We will first take the innermost — the transversalis — 
STRATUM of the ventral muscle, the stratum that lies internal to 
or beneath the osseous skeleton, or between the innermost parts 
of the skeleton, and internal to the chief nervous and vascular 
trunks. In most Fishes (though not in all, p. 807) it is absent, 
or partial or rudimentary, being represented by tendinous or 
fascial structures and scarcely distinguishable from the fascia 
tranaversalia. It may enter with that fascia into the for- 
mation of the post-cardiac diaphragm; and it is sometimes 
(Dog-fish) a main constituent of the pre-cardiac diaphragm. In 
Cryptobranch we have found it very extensively disposed beneath 
the ribs and the rest of the wall of the abdominal cavity, form- 
ing the depressores cosiarum and the transversalis muscle. It 
also lies beneath the vertebral column, from the pelvis to the 
head, forming the subvertebral rectus; and there is no dia- 
phragm. In higher animals it seldom is present as a continu- 
ous sheet in this last situation ; but portions of it remain, form- 
ing the longus colli, the rectus capitis anticus, the crura of the 
diaphragm and the retractor ani. On the interior of the lateral 
parts of the abdomen and thorax it is usually present in animals 
above Fishes, forming the transversalis muscle and the depres- 
sores costarum^, also the internal intercostals and the triangula- 
ris stemi* with the suhcostals. It is continued, with more 

^ These last are strongly developed in Snakes, and are quite segmented from 
the traTuvtrsalis with an interval between them and it, in whicli the Uvatore$ 
costarum appear. 

* The triangtilarii stemi and the tranivertalis are sometimes oontinnons in 
Man, as remarked by Stannius, Rosenmuller and Meckel. 



310 PROFESSOR HUMPHRY. 

or less interruption, within the pelvis and is then gathered in 
around the bowel as a pelvic diaphragm, o^;* levator ani, some of 
its fibres being reflected into, and some continued into, the 
muscular coat of the bowel In some instances — Fseudopus P. 
(p. 289), Cryptobranch (p. 17) and others — a part of it is con- 
tinued on into the subcaudal muscles. In Mammals a stratum is 
inflected from the ribs, or gathered in like the levator ant, upon 
the alimentary tube, forming the lateral portions of the post- 
cardiac diaphragm which, in continuity with a remnant of the 
subvertebral portion (the crura), coastitutes a septum shutting 
off the thoracic from the abdominal regions, but with apertures, 
like those in the pelvic diaphragm, for the passage of the oeso- 
phagus and the vascular and neural trunks. The psoas magnus 
and parvus also appear to be derivatives from this stratum ^ 
as well as some fibres of the crema,ster. 

It will be evident that the direction of the fibres of this 
stratum varies much in different parts of its extent, being antero- 
posterior where it lies under the vertebral column, oblique 
where it lies under or between the ribs, transverse where it 
forms the abdominal wall, and convei^ging where it is inflected 
upon the viscera; and in proportion to the diversity in the 
direction of their fibres the several parts are segmented from 
each other. 

In Cryptobranch it is throughout, or nearly, traversed by 
septa, and is so held in continuity with the transversalis Jascia, 
on the one side, and the rest of the ventral muscle and the com- 
ponents of the osseo-cartilaginous skeleton on the other. In 
higher animals, however, the septa for the most part disappear 
in its substance and upon its surfaces. They still abide forming 
the lines of separation between the digitations of the transver- 
salts muscle and the diaphragm, as well as between the mem- 
bers of the internal intercostal series, blended in the latter 
situation with the costal periosteum or perichondrium. 

The internal or transversalis fascia is in contact with the 
greater part of the extent of the internal surface of the trans- 
versalis stratum and is gathered in with it to form the levator ani 
and the post-cardiac diaphragm. Thus it lies upon the anterior 

> In the Omneft Pig the ptotu parvus passes with the p8. magnut and the 
iUaeue over the pnbes to the lesser troohanter of the femor. 



HUSCLEa IN VERTEBRATE ANIMALS. 311 

and posterior surfaces of both those diaphragms and reaches 
and is continued upon the alimentary tube\ From the 
anterior surface of the levator ani it is reflected upon the 
rectum and the bladder; and from the posterior surface it 
is continued to the integuments of the anus and the superficial 
^ fascia of the perineum. The part of the internal fascia behind 
the gathering-in of the levator ani forms the true pelvic fascia 
Uning the obturator muscle; and it is stretched beneath the 
pubes as the triangular ligament. The part of the internal 
fascia in front of the diaphragm lines the intercostals and the 
ribs and is gathered in, in front of the first rib, as the precardiac 
diaphragm, or the deep cervical fascia, between the thoracic 
and the cervical regions. The internal fascia is also continued 
upon the inner surface of the cremasteric projection of the 
transversalis muscle, forming what is called the fascia propria of 
the spermatic cord. Just in front of the pubes, in Man and some 
other animals, the internal fascia is separated from the tranaver- 
salis muscle which there passes with the internal oblique in 
front of the recites. 

The MIDDLE — the internal oblique — STRATUM is the most 
persistently intra-skeletal of the three strata of the ventral 
muscle ; and the intermuscular septa traversing it are, in many 
places, solidified by conversion into cartilage and bone. Thus, 
the ribs and sternum, the limb-girdles and the hyoid are deve- 
loped in it, and chiefly in its inner layers, though they may, and 
occasionally do, the limb-girdles more particularly, grow through 
the other strata and come into contact with the fascia transver- 
salia, on the one side, and with the skin, or the fascia superfid- 
alisy on the other. 

In the abdomen this stratum forms the interaal oblique, the 
quadratus lumborum^, the rectus* and the pyramidalis^. The 

^ Li Fishes these diaphragms are chiefly formed, so far as they are repre- 
sented at all, by the fascia alone. The sheet passing npon the bowel near 
the anns represented in Lepidosiren (Fig. 23) is the oiily representative of 
the pelvic diaphragm. 

The post-cardiac diaphragm is wanting in Sanrians as well as Ophidians 
(Stannins, 105). In some birds/ as Apteryz, it is well developed in its cmral, 
or vertebral, and costal parts, but is deficient in front, where the heart projects 
"jk through it. 

^ This muscle is carefully described by Gams, Beitrage zur vergleichend. 
Mufikellehre, Zeitschrift fiir ZoologUy III. He regards it as part of the ventro- 
lateral system ; he says it is absent in Birds, but present in ChelonianB, Ophl- 



312 PROFESSOR HUMPHRY. 

complete segmeDtation of the quadr, I. from the stratum, above, 
and of the rectus and pyramiddlia, beneath, is due to the sud- 
den alteration in the direction of their fibres from that of the 
fibres of the internal oblique. The relations of this muscle to 
both are nearly the same, a layer of it being continued upon the 
superficial, and another layer upon the deep surface of each ; 
and these layers, forming sheaths for the rectus and quadr, I,, 
axe in contact, and more or less blended, with the superficial 
(external oblique) and deep (transversalis) strata of the ventral 
muscle. 

In the thorax the middle stratum forms the series of exter-- 
nal intercostals. The direction of the slant of the intercostal 
muscles might indicate the external layer to belong rather to 
the superficial, or external oblique stratum and the internal 
intercostals to belong to this middle stratum. But, as we have 
abeady found, the fibres in different parts of the same stratum 
are disposed in very variable directions ; and the relations to 
the intercostal nerves and blood-vessels are stronger grounds for 
classing the external intercostals with the internal oblique and 

dians, Samians and Batrachians; that in Man it sometimes passes to the body 
of the 11th dorsal vert, and to the 11th rib, in the Buffalo to the 4th hinder dorsid 
Tert. and ribs. In some animals, as Hyrax (see Meckel), it advances still farther 
forwards. In Cryptobranch its representative is not segmented from the rest of 
the ventral muscle, but, like that of the rectus, is continuous with it. In 
Pseudopus Pallasii it is distinctly a serial correspondent and extension of the 
intercostals passing from the hincUnost rib to the ikac bone (p. 289). 

' The relations of the recttis to the several strata of the ventral muscle vary 
a good deal. Primitively, as in Lepidosiren and most Fishes, it is continuous, 
and on a level, with the whole thickness of the ventral muscle. In Cryptobranch 
it is so with the middle and external strata only, while the internal stratum 
passes above it. In Snakes a deep part of it seems to be represented by 
thickening of the intercostal fibres near the extremities of the ribs. Some of 
these thickened bundles pass from rib to rib, and some from one rib to another 
at a greater distance. And in them, and more clearly in Pseudopus P. (p. 288), it, 
or the chief part of it, lies on a plane superficial even to the external stratum 
the fibres of which pass into its deeper surface. In Mammals it is completely 
segmented from all the strata, and lies in the greater part of its course between 
the layers of the middle stratum. 

Miiller, AbhandL der Berlin. Akad, 1834, p. 345, and Tab. I, describes and 
represents the rectus as distinct from, and covered superficially by, the obliqvi 
in Bdellostoma. This is, however, unusual in Fishes. 

* The pyramidalis appears to have special relations to the marsupial bone, 
and to the spine and crest of the pubes, which are the representatives of that 
bone ; and its presence as an independent muscle is probably to be associated 
with the tendency to the presence of this as an independent bone (footnote on 
p. 16). Functionally, it seems to assist in strengthening the suprapubic region 
of the abdominal wall; and it is well developed in animals ffemsJe Pteropus, 
Journal Anat. itl 301, and Pseudopus P., on page 289) in whicn, in consequence 
of an interval between the pubic bones, the recti muscles diverge and leave this 
part of the abdomen comparatively unsupported. 



MUSCLES m VERTEBRATE ANIMALS. 313 

the internal intercostals with the transversalis stratum. — ^The 
levatorea costarum which, like the external intercostals, are 
overlapped by the dorsal muscle, may also be ranged as mem- 
bers of the middle ventral stratum \ 

The rectus is usually continued forwards on the external 
surface of the sternum and costal cartilages, not unfrequently 
as far as the first costal cartilage, constituting a recttis thoracis. 
Other longitudinal muscles are sometimes (Ai, Cyclothurus, 
Otter, &c.) developed, lying also beneath (superficial to) JJie ribs, 
more laterally than the rectus, and constituting what may be 
designated recti thoracis laterales^y to distinguish them from, 
and mark their relation to, the rectus thoracis. These muscles 
approach or alternate with extensions of the scaleni back- 
wards. The rectus may (Cyclothurus and Otter) be crossed 
superficially by a stemo-costal muscle passing from the sternum, 
obliquely forwards and outwards, to the ribs and approaching 
the scaleni. All these belong to the middle or internal stratum 
of the ventral muscle and are merely illustrations of varying 
cleavage and segmentation in accordance with varying direc-> 
tion of fibres. 

From the ribs the middle stratum is continued forwards 
laterally to the cervical transverse processes as scaleni. These 
are separated from the rectus capitis and longus colli of the 
internal stratum (p. 309) by the cervical nerves. As just hinted, 
they often extend upon the exterior of the ribs, meeting the recti 
th. laterales and the stemO'Costales*, Anterior to the scaleni is 
the recttts capitis lateralis. Nearer the mesial line, inferiorly, 
the middle stratum is continued forwards to the hyoid*, thyroid, 



^ In Snakes, these last are much deyeloped and extend over a greater range 
than in other animals, for the purpose of assisting in progresdon ; and there is 
oommonly to be found a series of internal Uvatores coitarwn, situated external to 
the outer part of the depressores, crossing those musoles and appearing in the 
interval between them and the iraruvenalU, Their position would indicate 
them to belong to the internal stratum ; but the nerves which lie external to 
the depressors and the transversalis, pass internal to the levators. It may be 
added that the nerves in these n-nimAlw do not confine themselves to the intercostal 
spaces opposite which they escape from the vertebral canal, but, in some 
instances, cross over one or more ribs, passing between the ribs and the trans- 
versalis. 

' One of these has been so named by Macalister, in his description of AI, 
AnnaU and Mag. of Nat. Hut., June 1869. 

> In the Oreen Monkey the seaUntu is continued into the reetitt. 

* The inscriptions in the ventral muscle running forward to the hyoid. 



314 PROFESSOR HUMPHRY. 

tongue, jaw and pharynx as stemo-hyoid and stemo-ihyroid, 
as hyo-glo88U8 and genio-hyo-glo8SU8, and as hyo- and thyro- 
pharyngeus, or middle and inferior constrictors of the pharynx. 
These come into relation with muscles ascending to, or de- 
scending from, the skull, which probably also belong to the 
same stratum, viz. stylo-glossua, stylo-hyoideua with the hinder 
portion of the digastricus^, atylo^haryngeus, superior constrictor 
of the pharynx, as well as the faucial and palatal muscles. 

From the outer surface of the middle stratum muscles are 
detached to the shoulder-girdle. Those passing to the part 
of the girdle above the glenoid cavity (the scapular part) con- 
stitute the costO'Scapular or serratus group. They consist of 
one or more muscles attached to the ribs, interdigitating and 
•sometimes connected by continuity of fibres with the external 
oblique, so as to present strong claims to be regarded as part of 
the same stratum with it ; but in their course they are deep ; 
and they are inserted into the deeper surface of the margin of 
the scapula, on one or two sides, between the sub-scapvlaris and 
the other muscles which are inserted into, or near, the margin 
of the scapula. Anteriorly, this group is prolonged into the 
neck as the levator scapuks which is not unfrequently conti- 
nuous with the serratus, and as the omohyoid^ The muscular 

present in Lepidosiren and others, are not nnfreqaently represented by one 
inscription near the middle of the stemo^hyoid. This inscription is continued 
into the omohyoid with which the stemo-hyoid is sometimes continuous ; and 
it is usually persistent in the latter muscle, in Man, where it makes a bend 
towards the clayiole. Stannius speaks of an extension of the rectus to the tongue 
in Triton. 

^ In the Porpoise this part of the digastric only is present passing from the 
hinder part of the temponil to the hyoid. It is called occipito-hyoid by Bapp, 
Die Cetaeeen Zoologiaeh-anaUnnisch dargesteUt, s. 182, and by Stannius, MiUUr's 
Archiv, 1849, s. 7, but is regarded by Stannius as the posterior beUy of the 
digastric. For account of the anterior part see page 824. 

* The omohyoid may be, as in Phooa, continuous with the atemo-huoid^ 
forming a broaa musde inserted into the sternum, the ulnar tubercle oi the 
humerus and a fascial band between the two. It may, as in Sdno, be attached 
to the claTicle as weU as to the scapula ; and the ttemo'kyoid is in that animal 
attached to the InterclaTiole, or epistemum. Tendinous traces of the claTi- 
oular attachment bend it towards the clavicle in Man, and are, partly or entirely, 
the cause of the persistence of the inscription or interruption in its musenlar 
fibres. Though m the same plane with the levator scapula, it is separated from 
it by the situation of the branchial opening, which also separates the members 
of the superficial brachio-oephalio stratum (the stemo-mastoid and trapeaius) 
that overlie this region. In the Hippopotamus it passes from the side of the 
basi-hyal and the sub-hyoidean septum to the undw surfiMe of the oooipito- 
humeral part of the trapesius which it joins at an angle, and is united to it by 
an inscription. The foremost fibres run on without any definite inscription to 



1 



n 



MUSCLES IN VERTEBRATE ANIMALS. 315 

derivations from this layer, which pass to the girdle beneath the 
glenoid cavity, are the stemo- or costo-coracoids. When the 
coracoid is abortive, the costo-clavicular, called subclavius, may 
take the place of the costo-coracoid^ ; and when both coracoid 
and clavicle are abortive, the member or members of the group 
may pass to the under surface of the scapula, constituting the 
costo- or 8temo'Scapulars\ Thus, spreading beyond their usual 
limits, these may come into close relation with the serratus 
group. Or the serratus, extending lower down than usual, 
may come into close relation with the costo-coracoids. Still 
the two groups — the costo-scapular and the costo-coracoid or 
costo-clavicular — are, I believe, always separated by the nerves 
to the limb passing between them. 

These two groups of muscles passing, respectively, to the 
scapular and to the coracoid or the clavicular parts of the 
shoulder-girdle, together with the muscles passing forwards to 
the hyoid, tongue and jaw, constitute that which I have desig- 
nated (pp. 262, 285) the deep brachio-cephalic stratum of 
the VENTRAL MUSCLE. In animals (Snakes and Uro- 
^ delans), in which the inferior wall of the thorax is not closed in 

V by the sternum and the costal cartilages, the middle, or cepha- 

lic, part of the stratum may be continued onwards, uninter- 
ruptedly, to the hyoid and the mandible, but it is not seg- 
mented from the superficial stratum (Fig. 44 and description) ; 
or (Fishes, Figs. 26 and 38) it may be interrupted by the cora- 
coids extending athwart the middle line and uniting with one 
another, and may then be segmented from the superficial stratum. 
Traced backwards from the abdomen, the middle stratum 

the htunems. In the Pig it passes over the anterior edge of the scapula to the 
upper edge, and is connected by fibrous tissue with the radial tubercle of the 
humerus and the deltoid muscles. 

1 These two do not coexist, when the coracoid is large, the muscle passes 
from the sternum or rib to it, and forms a itemo^ or costo-coracoid. When the 
coracoid is short the muscle, if present, passes from the rib, or the first or 
(Orycteropus) second costal cartilage and perhaps the sternum, to the claTicle 
oonstituting the nibclaviui. It may, as it does in Oiycteropus and Seine, 
extend on to Uie scapula. In the Porpoise the eiibelaviui passes from the first 
rib, near the sternum, to the coracoid; and the pect. minor passes from the 
second rib to tiie humerus. Stannius, "Beschriebung der Muskeln des 
Tilmmlers.'* MUller's Archiv, 1849, s. 14 and 16, caUs the former of iheBepect, 

\^% minor, and the latter eosto-kuTneralis. 

\ * The stemo-scapular may coexist with the subclavius. Thus Maoalister 

found the subolaTius quite separate from the stemo-scapular and iiuerted into 
the claTicle. In such case the muscle is divided into two. 



31 G PROFESSOR HUMPHRT. 

of the ventral muscle encounters the pelvis and is interrupted . 

by it*. Its continuations backwards from the pelvis constitute 
the iliO' and tschio-caiuiales*, the deep sphincter-ani, the erector- 
penis, compressor-urethrce and transversus-perinet, besides the 
extensions upon the hind limb around the hip-joint to be sub- 
sequently mentioned. ^ 

I pass now to the disposition of the external stratum 
of the VENTRAL MUSCLE which has the external oblique 
muscle of the abdomen as its most steady representative, and 
which I, therefore, sometimes call the EXTERNAL oblique 
STRATUM. As already stated, it is the stratum in which the 
ossifications, not preceded by cartilage, most frequently occur. 
The 'membrane bones' thus formed are commonly in the 
situation of the septa, and, therefore, overlie the 'cartilage 
bones' formed in the septa of the middle stratum. Some- 
times they are blended with them, the two being ossified toge- 
ther, or the one may serve as a substitute for the other. Thus 
the epicostals', the episternum^ and the epicoracoid, or clavicle, 

^ It mast not be forgotten, that the rudimentaiy peine bones of Ophidians 
lie, or rather project upwards, internal to the transversalU. Perhaps they ^^^ 

originate in the internal oblique stratum, and grow through the transyersaUs, 
just as in many animals the ilium presents through the external oblique 
stratum, and as in the Dog-fish, the scapula pushes its way into the dorsal 
musole (Fig. 28). In Pseudopus P., the ilium, though deep, is not quite so deep 
as in Sn^es, a considerable part of tiie ventral muscle is inserted into, or 
interrupted, by it, and a distinct strip of the muscle passes beneath it (p. 289). 

' The ischio-caudal, and the ilio-caudal are, to some extent, serially homo- 
logous, respectively, wilii the costo-coracoid, and the oosto-scapular {terratus m.); 
and the nerves to the hind limb pass between them, as do the nerves to the fore 
limb between the last-named muscles. 

The shoulder-girdle, like the pelvic girdle, varies in the depth at which it 
is placed, and also remarkably in the size of its eoraeoidal part, and in its 
connection with the rest of the skdeton above and below. In Fishes the 
ooraooid part is imbedded in the deep stratum of the ventral muscle ; but in 
Ciyptobranoh, the coracoid, notwithstanding its size, has little direct connection 
with the ventral muscle, the deep layers of whidb pass clear of it and above It. 
In Birds and Reptiles its connection with the deep ventral stratum is re* 
established by the costo-coracoid muscles. In MftTmrmlg the connection is 
chiefly, through the medium of Uie peetoraHs minora with a deep layer of the 
pectoral or superficial stratum of the ven^aJ muscle. 

' In Hatteria the epioostals are more numerous than the costals, occupying 
not only the lines over them, but the interspaces between those lines : see Giin- 
ther, PML Trans. 1867, p. 608, whose description I have in most points verified. 

^ I much prefer the old term, * epistemum ' to that of 'interdavicle*, proposed 
by Mr Parker ; because it expresses not only its position, but its nature as a bone 
fonned in the membranous tissue upon the sternum, at the same time that it 
refers it to the same series as the epioostals and the clavicle. — ^It will be per- 
ceived that, for the same reason, I use the term ' epicoracoid' to designate, not 
the cartilage or cartilages lying between the coracoid and the sternum, but tbo 



MUSCLBS IN VERTEBRATE ANlMALa 317 

\ are formed, respectively, over the costals, the sternum and the 

I coracoid, and the lower jaw is formed over MeckeFs cartilage. 

We have seen that this stratum is, in Fishes, closely connected 
with the skin by the transverse and longitudinal septa passing 
through the compact intervening external fascia. 

A superficial layer of the stratum is often, more or less com- 
pletely, segmented from the rest, and, retaining its connection 
with the skin, or with the superficial fascia or both, constitutes 
cutaneous or subcutaneous muscles to which the names pan- 
nicle, platyama myoides, &c. are given. They may retain their 
original connection not only with the rest of the external ven- 
tral stratum, but also with the deeper strata, and with the car- 
tilage bones. Thus they, in some instances, are united to the 
ribs and the sternum, the vertebral spines and the limb-bones. 
They often correspond, and are more or less blended, with the 
several divisions of the rest of the stratum, and will therefore 
be best considered in relation with them. They commonly ex- 
tend over the lateral septum, spreading upon the dorsal muscle, 
and reaching the dorsal mesial line. This also we shall find to 
be the case with the rest of the stratum. Or, which is much 
\^^ the same thing, the components of this external stratum of the . 

ventral part of the lateral muscle are confluent with superficial 
dorsal fibres, and form one stratum with them; so that it 
might be called the ' external ventro-dorsal', or, better, the ' ex- 
ternal lateral' stratum, that is, the external stratum of the 
entire lateral muscle \ See Diamond Snake (Fig. 44)'. 

membrane bone, or olaviole, formed upon it, and often in doee connection 
with it. 

^ This extension of the external stratum of the yentral part of the lateral 
mnsde, oyer the lateral line and over the dorsaJ part of the musole^ to the ver- 
tebral spines is due, perhaps, to the expansion of the visoeral cavity, laterally 
and upwards, as seen in the section of the Tadpole (Figs. 42 and 43). 

' Description of Fig. 44, PI. xvi. — The Diamond Snaxe. 

Dissection of the fore part of a Diamond Snake. — The pin is inserted into 
the fore part of the oerato-hyoid cartilage which is seen ronning badt among tile 
muscles. The drawing shews the bundles of the external oblique arising partly 
from the lateral septum, and partly from the fascial tissue on the surface of the 
dorsal musde and passing, downwards and backwards, into the antero-pos- 
teriorly directed fibres of (r.j the rectus abdominis. Traced forwards, the rectus 
is seen giving off the bundles of the external oblique. Anteriorly, it expands 
into a muscular sheet, in the middle of the thickness of which the hyoid is in- 
volved. This runs forwards to the lower jaw, covers the under part and sides of 
the throat, expands upon the side of the head and neck, and extends. over the 
dorsal muscle to the cervical spines. A set of its bundles more distinct than 
«. the others runs to the angle of the mouth, constituting a retractor oris. 

This expansion is a continuation of the entire thickness of the ventral mus- 

VOL. VL 22 



318 PROFESSOR HUMPHRY. 

. In the abdomen the stratum is chiefly represented by the 
external oblique which extends upon the exterior of the thorax, 
and the fibres of which often blend with the fibrous tissue over 
the dorsal muscle. Anteriorly, it is continued, or its superficial 
fibres are continued, with a variable amount of interruption, 
into a 'superficial brachio-cephalic' sheet which extends to the 
face and head, which is distorted by encountering the fore limb 
and its girdle, and the fibres of which converge upon the limb, 
as though, like the skin, they had been pushed before it as it 
grew out from its girdle. Hence, opposite the limb, the fibres 
are directed transversely, from the ventral and dorsal mesial 
lines, upon the limb; whereas, before and behind, they have a 
more oblique or antero-posterior direction. 

As it advances towards the fore limb the inferior, or ventral, 
part of the superficial brachio-cephalic sheet resolves itself into 
the pectoralis major, the fibres of which converge upon the ra- 
dial edge of the humerus, inclining to the plantar aspect ; the 
superior, or dorsal, part of the sheet resolves itself into the latis- 
simus dorsi, the fibres of which converge upon the ulnar side of 
the humerus, and are often, to some extent, blended with the 
muscles on the dorsal aspect of the limb. I say, to some ex- 
tent, because the prolongations upon the middle and distal 
segments of the limb of both the inferior and superior parts of 
the sheet shew an inclination to the plantar aspect, which 
is a consequence of the position and flexures of the limb. The 
ivro^-pectoralis and lat d. — may (Lepidosiren, p. 259) be united 
and pass as a continuous sheet upon the radial and ulnar mar- 
gins and upon the intermediate, plantar or axillary, that is, the 
posterior aspect, of the shoulder and limb. More commonly 
they are separated by the axillary interval; or, connecting axil- 



ole, and oontains therefore the elements of all the mnscles coyering the under 
part and sides of the neck, and passing between the thorax, hyoid and jaw. It 
represents, in short, the whole of the braohio-cephahc stratam of Lepidosiren 
and Dog-fish, — the cervicalis profundus, that is, as well as the ceryicaUs snper- 
ficialis ; — and it includes therefore the factors of the stemo-mastoid and trape- 
zius, the platjsma, constrictor faudum, <fec. A continuation of it backwards 
would represent the pectorals and latissimus dorsi as well as the serratus. 

Though its hinder edge seems to be marked off from the oblique, yet its con- 
nection, and the connection of the oblique with t^e rectus, and the fact that the 
oblique bundles cease where this more continuous expansion begins, shew that 
it is a serial continuation of them, of the parts of them more especially which 
extend upon the dorsal muscles. 



MUSCLES IN VERTEBRATE ANIMALS. 319 

lary bands, may remain as a result of imperfect segmentation*. 
^ The term 'achselbogen' has been applied to them. The term 

'axiUary* is that by which I will designate them. 

The cosio-alaris {costo-anconeua) of the Bird is an interesting 
y example of one of these * axillary * muscles, being segmented from 
both the pectoral and the latissimua dor si, and passing from the ribs, 
between the two, near to the aerratus^ along the inner side of the arm, 
to or near the inner condyle of the humerus. In the Swan I found 
part of it elastic, where it occupied the hinder fold of the axilla, and 
traced it along the ulnar margin of the wing, and in the retiring angle 
beneath the carpus, to the skin and tissue binding the skin to the 
ulnar phalanges. Thus it presents interesting antagonistic homo- 
logical affinities to the tenMor pliccB cdaris ; and by its near relation to 
the serroUus on the deeper side, and by its blending with the cutaneous 
muscles and the skin on the superficial side, it is an interesting relic 
of the primitive union of the several strata. 

The relation of this brachial, or hinder part, of the superficial 
brachio-cephalic sheet to the rest of the external oblique stratum 
varies a good deal. In many animals, for instance, the pectoral is a 
continuation of the whole thickness of the external obliqtie, or of the 
whole or great part of the rectus. In Cryptobranch it is almost 
entirely derived from these. In others, it is a superficial stratum 
only of the oblique, extending perhaps to the pubes. In others again 
it arises from the outer surface of the aponeurosis of the obUque ; 
and in Hatteria it appears to arise from between its layers'* In Man 
it is usually continuous with the obliqiie by means of only a few 
muscular fibres. In the Hippopotamus it is largely continuous with 
the subcutaneous muscle over the oblique ; whereas in Birds its 
origin is so confined to the sternum and the clavicle that we should 
little have thought of regarding it as an extension of the abdominal 
stratum, had our attention been limited to its anatomy in them. In 
like manner the Uuiasimus dorsi is, in some, confined to the neural 
spines, in others extends upon the ilium; and, not unfrequently, it 
derives origin from the hinder ribs, where it may (Seal) meet, and be 
united with, the pectoral. In the Porpoise its origin is confined to 
the 5th, 6th and 7th ribs'. 

^ A good ninstration of these axiUary masoles was presented this winter by a 
female subject in the disBecting-room of this University. A slip from the lower 
edge of the peetoralUt and another from the anterior edge of the latisHmtu 
dorsi in eaoh arm, ran through the axiUary space and, meeting, were inserted 
together into the fascia ooyering the coraeo-braehialU. 

These connecting bands are commonly large in Camivora. They are osnaUy 
formed by an extension of the latUtimus dorsU or parts of it, beneath the axilla, 
to the pectoral muscle and the pectoral ridge of the hnmerus ; while another 
part of the lat. d. runs along the inner side of the triceps to the inner condyle 
forming the dorH-epitrocMien, 

• Phil. Trans. 1867, p. 609. In Pteropns, Journ, Anat, iii. 300, it is in 
three separate piurts, an abdominal, a sternal, and a clavicular. 

' The variation in the mode in which cleavage of strata may take place is 
illustrated by the fact, that in Birds and Reptiles the lat. d, usually overlaps the 

22—2 



S20 PROFESSOR HUMPHRY. 

All this hinder, post-brachial, portion of the external stra- 
tum of the lateral muscle may be covered by a superficial, or 
subcutaneous, layer, which varies much in thickness, and which 
may be connected with one or all of the divisions of the stra- 
tum, sometimes reciprocating with them in size and thickness, 
as in the Hippopotamus. Like them it converges upon the 
arm ; and, accompanying the pectoral more particularly, it may 
be lost in that muscle, or it may be inserted into the humerus 
or the coracoid. 

It is often a muscle of such size, and of so powerful action upon 
the arm, and radiates from the arm so widely upon the flank, that 
I have^ described and named it as brachio-lcUeral, Sometimes it, or 
part of it, lies upon and is closely \mited with the laJtissimua dorsi; 
or it may be (Hatteria) continuous with its lower edge. Sometimes 
(Manis) it is difficult to separate from the skin on the one side and 
the external oblique on the other '. As mentioned above, it forms part 
of the cosUHdaris of the Bird. 

Traced forwards into the pre-brachial or cervical region the 
SUPERFICIAL BRACHIO-CEPHALIC STRATUM, which we are con- 
sidering, is, after a certain interruption caused, partly, by the 
shoulder-girdle, and, partly, by segmentation from alteration of 
the direction of its fibres, continued as a superficial cervical sheet 
^-cervicalis superfidalis. It covers the neck, from the shoulder- 
girdle to the skull, and from the ventral median line to the 

trapezius^ the reverse being the case in Mammals. In the Rabbit the two mus- 
cles are continaons. 

^ Joum, Anat, iv. 27. 

* The superficial, cutaneous layer is well developed in the Porpoise, where 
it spreads upon the fins, under the abdomen, and under the sheath of the penis ; 
and in the female it passes beneath the mammary gland, detaching fibres which 
encircle the lower dilated part of the duct, so as to compress it during suckling. 
It attains its maximum in the Hedgehog, being connected with the Cental and 
occipital bones, the lower jaw, sternum and humerus, and the dorsal and caudal 
spines, and forms a thick sphincter around the body beneath the line where the 
bristles terminate. 

In the Guinea Pig both pectoral and trapezius are continuous with the pan- 
nide. In the Pig, and many Animals, the brachio-lateral part extends over tLe 
thigh. In the Hippopotamus it is veiy largely developed, forming a thick mass 
over the abdominsd muscles, which are idmost dwajfed or supplanted by it, 
though quite distinct from it ; it extends over the thigh and the Imee ; and it is 
also continued into the pectoral. 

In Snakes the continuity of strata is manifested and maintained by the 
bundles of the oblique abdominal muscles passing from the ribs direcUy into the 
ventral scutes, which are thus enabled to act as organs of locomotion. Some of 
the bundles running horizontally for a distance, from scute to scute, constitute 
(pp. 818, 827) a superficial or subcutaneous reettu. 



iniSCLES IN VERTEBBATE ANIMALS. 321 

lateral line ; and it commonly extends, in a muscular form, be* 
yond the lateral line, to the dorsal median line. Anteriorly, it 
is connected with the whole circumference of the skull, and 
sends superficial prolongations over it. It is well exhibited in 
Lepidosiren (Figs. 24 and 26), Dog-fish (Figs. 28 and 29), and 
Ceratodus (Figs. 34 and 37), and I have named the ventral 
part of it, lying beneath the level of the gills, cervicalU super- 
ficialis inferior (C 8, i.), and the dorsal part of it, lying above 
the level of the gills, cervicalia superficialis superior (C 8. 8.). 
These two parts are, however, continuous upon and above the 
gill-cover; and the superior or dorsal portion {C. 8. 8.) seems 
rather to be an extension from the inferior or ventral portion 
(G. 8. t.), just as the ventral and dorsal portions of the post* 
brachial part of the same stratum are often continuous, and the 
latter seems to be an extension from the former. 

In animals like those just mentioned, where there are 
gills with the attendant opercular structures, the cervicalia 
superficialis is but a thin expansion, and does not present 
any distinct segmentation beyond that— often not very dis- 
tinct, into levatores, depressores, &c. arcuum — ^which is requisite 
for the movements of the branchial apparatus. Where, how- 
ever, the gills are abortive it acquires increased thickness and 
is stratified and segmented. 

A superficial layer forms the subcutaneus colli or platys^ 
ma which' covers the neck and is prolonged forwards upon 
the fstce and head, where it blends with and is continued 
into the facial muscles, the auricular muscles and the ocdpito- 
frontalis. It is prolonged backwards upon the thorax and 
fore limb to an indefinite extent, blending with the bra^ihuh 
lateral and, like it, finding its way to the osseous structures 
of the limb. As in the case of the brachio-laterai, portions of 
it may blend with, or supplant, segments of the subjacent 
layer. 

In some instances it is continuous with the trapeziue; and in 
Pteropus it takes the place of the cervical portion of that muscled 
A remarkable detachment firom between its dorsal and its ventral 

^ Maoalister finds that the miiBole, which thus in Bats supplants the cervical 
part of the trapexitu, is supplied not by the spinal accessory, bat hj cervical 
nerves and the seventh nerve. 



322 PROFESSOR HUMPHRY. 

portions, blended with a similar detachment from the deeper layer 
(from the contiguous edges of the pectoral and trapezio-deltoid), 
jfbrms the tensor plicce alaris, better called cervic<halari8. This, partly 
composed of elastic tis>^ue, extends along the radial edge of the wing 
in Bats and Birds, and reaches the skin and subcutaneous tissue of 
the ituiial digits. It ia thus (as mentioned p. 319) the antagonistic 
homologue of the coato-alaris. 

The deeper layer of the cervicalis superficialia developes 
into the atemo-cleido'mastoid and the trapezius, the former 
representing the ventral, or sub-branchial, part of the layer, 
and the latter representing the dorsal, or supra-branchial 
part*. They are very regular muscles, but vary in their 
range of attachment at both ends. The stemo-cleido-mastoid 
is sometimes confined to the sternum and then is called 
sterno-mastoid. Sometimes it is inserted, by a narrow tendon 
only, into the mastoid, instead of by a broad musculo-tendi- 
nous termination as in Man". Sometimes the portion con- 
nected with the clavicle is separate, forming a cleido-occipital. 
The trapezius, instead of the extensive origin from skull, 
cervical and dorsal spines, overlapping the latissimus dorsi, 
which it has in Man, may (Pteropus and Birds) have much 
more limited origin. It may, in the absence of the clavicle, 
be inserted (Cryptobranch) into the anterior edge of the scapula 
and the precoracoid or (Ai) the coracoid; or it may be conti- 
nued as trapezio-deltoid to the humerus*. When this last 
occurs, there is usually an inscription remaining between the 
trapezius and the deltoid, in the deeper part of which a 
rudimentary clavicle is often present in Camivora. 

From beneath the trapezius the rhomboids* are segmented, 

^ The interval between the two in Racoon is partly bridged over by fibres 
passing from the trapezius to the sterno-mastoid, reminding ns of the epi- 
branohial fibres connecting the superior and inferior cerviccUea parts in Lepi- 
doslren and Dog-fish. 

' In a Fawn I found it extending over the lower jaw to the orbital edge of the 
maxilla, occupying that part of the area of the superficial layer. In Crocodile 
it passes beneath the trapezius to the transyerse process of the 4th cervical ver- 
tebra, where it comes into contact with the middle ventral stratum represented 
by the Uvator scapula, 

* In Manis it extends over the biceps and brachial vessels to the internal 
condyle of the humerus. 

* The rhomboid in Owl and Kite extends from all the lumbar and dorsal 
spines to the hinder edge of the scapula, dwarfing the trapezius. In Apteryz it 
U said by Owen to be wanting. It is present in the Crocodile, but I have not 
met with it in any lower animal; sometimes it blends with the latissimus dorsi. 
Joum, Anat. iv. 83. 



MUSCLES IN VERTEBRATE ANIMALS. 323 

also apparently the serrati postici, as well as the masUhhume^ 
ral and Tnasfxhscapular. Though the last two might perhaps 
be said to be segmented from the deep surface of the stemo- 
mastoid, yet they usually blend below with the trapezius 
or deltoid, or pass to the scapula or humerus. Also another 
muscle is segmented from the trapezius, which I have in 
foimer papers named cervico-humeral, and which is an in- 
teresting reminder of the fact that in Lepidosiren (p. 260) 
and Dog-fish (p. 274) some of the deeper fibres of the layer 
are traceable to the lateral septum and, beneath it, to the 
lateral parts of the vertebrae; for the cervico-hwrneral arises, 
not like the trapezius from the vertebral spines, but from 
the transverse processes of the atlas or other cervical verte- 
brae, near to the levator scapulce, or even (Hedgehog) from 
the fore part of the ring of the atlas, or (Guinea Pig and 
Rabbit) from the basi-occipital. It descends upon the shoulder 
to the clavicle, acromion, or humerus, or blends with the 
trapezius or deltoid. Thus, though it is associated with the 
levator scapuke in its origin, it is associated with the trapezius 
in the latter part of its course; and it is evidently a repre- 
sentative, in Mammals, of those fibres of the stratum which, 
in Lepidosiren, detach themselves from the rest of the stra- 
tum and bend in, deeply, to the sides of the vertebrae. In 
the Hippopotamus the cervico-humeral arises by a round 
tendon from the back of the exoccipital, behind the digas- 
tric and stylo-hyoid, joins the trapezius, forming its fore part, 
and runs on to the deltoid ridge of the humerus. It is not 
separated from the deltoidal, or lower, part by any inscription. 
It is joined by the omo-hyoid which loses itself in this muscle. 
It is called masto-humeral by Gratiolet. A separate large por- 
tion arises by a strong tendon from the under surface of the 
transverse process of the atlas and spreads upon the dorsum 
of the scapula. The trapezius passes over the large trans- 
verse process of this vertebra without deriving any fibres 
from it. 

Between the two layers of the cervical part of the super- 
ficial brachio-cephalic stratum, just described — ^the superficial 
layer which is forming the suhcutaneus coUi, and the deeper 
layer which forms the stemo-mastoid and irope-nw— other 



824 PROFESSOR HUMPHRY. 

muscles are developed in different animals, to which the names 
cervid svimaodllaris, depressor mandibiUcB, myh-hyddeus and 
genio'hyoideus are given. These vary a good deal; and the first 
two are absent in higher animals. I have said that they are found 
between the two layers; but the genio-hyoid, which lies near the 
middle line, is perhaps rather to be regarded as segmented from 
the deeper surface of the superficial layer. The most remarkable 
member of this series is the anterior belly of the digastric. It 
lies in a more superficial plane than any, yet is continued, 
through the medium of a tendon or inscription, into the pos- 
terior belly which runs, in company with the stylo-hyoid, be- 
neath the stemo-mastoid, to a deeply situated spot of the tem- 
poral bone, just behind the styloid process. — The peculiar confor- 
mation and disposition of this muscle in Man, and the inscrip- 
tion which in many lower animals is substituted for the 
tendon between its two muscular portions, have excited much 
attention and been the cause of many surmises. I believe 
the real explanation of it to be that it is composed of two 
muscles or muscular portions derived from the two strata of 
the brachio-cephalic muscle. The hinder portion is, together 
with the stylo-hyoid, a derivative from the deep stratum, *^ 

which it will be remembered is an extension of the middle 
or internal oblique stratum of the ventral muscle; and the 
anterior portion is a derivative from the fibres of the cervi- 
calls superficialis layer, which lies next above the subcutaneus 
colli. It is an instance of a portion of a deep stratum re- 
taining continuity through, the medium of a septum with a 
portion of a superficial stratum. The septal remnant, which 
is a remnant of the hyoidean, or sub-hyoidean, septum (the 
septum, that is, in which the hyoid is formed), is the tendon 
of connection between the two portions and the band which 
holds them to the hyoid\ 

^ It may be observed, as affording some confirmation of the relation de- 
scribed in the text between the anterior belly of the digastiic and the mylo-hyoid, 
that they are both supplied by the mylo-hyoid branch of the 6th nerve ; whereas 
the posterior beUy of the digastric and the stylo-hyoid are both supplied by the 
7th nerve. I do not however attach much importimce to this. 

In this derivation of the digastric from the elements of deep and superficial « 

strato, we are reminded of the same thing in the cervico-humeral just mentioned, 
and in the eo$to-alarU (p. 819). 



MUSCLES IN VERTEBRATE AIOMALS. 825 

The dlsBection of the Hippopotamns* (Figs. 45 and 46) affords an 
interesting illustration and confirmation of these views. The sitbcu- 
taneu8 colli {S. e.) under the neck is of great thickness, as is the 
case with the subcutaneous muscles on the under sui-face of the body 
geuerallj ; and it api^ears at first sight to terminate in front in a thick 
defined border extending, on either side, upon the middle of the 
large mandibular protuberance into which the masseter (M) is in- 
serted. The muscle at and near this border is further thickened by 
transverse fibres crossing from one side of the face, over the mandibu- 
lar protuberance, to the other side. Close examination, however, 
shews that the anterior border is not so defined as at first sight 
appears; for the edge is connected by fibrous tissue, which is the sub- 
hyoidean septum', with deeper-lying muscle, a broad, thick hyo-menr 
tal (A, M.), which passes forwards, from the septum just mentioned, 
to the mandible on the side of the bulging symphysis. By means 
of this septum the hyo-mental is continuous, behind, with the super- 
ficial fibres of the stemo-hyoid {St, U,) which is a large thick muscle, 
on its deeper suri^u^e, with the basi-hyoid bone, and, on its superficial 
surface, with the edge of the subciUaneus coUi\ Above it, and deeper 
than it, is the mylo-hyoid {M, H,) Above this is (G. H,) the genio- 
hyoid j and deeper still is the genio-hyo-glosstis. These are disposed 
as they usually are in Mammals. The atyh-hyoid aiises, tendinous, 
from the outer side of the tooth-like exoccipital process, soon expands 
into a muscular belly which is, partly, inserted into the byoid, on a 

^ Description of Figures of the Hippopotamus and the Cat (PI. xvi.). 

Figs. 45 and 46. Dissections of the muscles beneath the floor of the month 
of a Hippopotamus which died three days after birth. — I.M.^ the inferior 
maxilla, with its large submental protuberance uncoyered. — L, the upper lip.— 
5. c, snbcntaneus colli. — ilf, Masseter. — U, M,, hyo-mental entire on the left 
side, but divided on the right side, and, in Fig. 46, reflected. — D,, anterior part 
of digastric running parallel with, and blended near the jaw with, the hyo- 
mental. — M. H., mylo-hyoid divided and partiaUy removed to expose G. H., the 
genio-hyoid. — F,A,,ihe facial artery.— In Fig. 46, the suboutaneos colli has 
been removed, exposing the hinder part {D' ) of the digastric, also S, H. , the 
stylo-hyoid which is seen to be continuous with the hyo-mental. — St, M, , sterno- 
mastoid. 

Fig. 47. — Similar dissection in Cat. S. c, the snbcntaneus colli, or platysma, 
has been divided and pulled aside to shew (Z> and D') the anterior and posterior 
parts of the digastric, with the inscription between them which is the sub- 
hyoidean septum. The anterior part of the digastric is seen to occupy the posi- 
tion occupied by the hyo-mentaJ, as well as by the anterior part of the di^^tric, 
in the Hippopotamus. — M. H., the mylo-hyoid. — St. fl"., stemo-hyoid. 

* It is called by Oratiolet, ItechercheM $ur VAnatomie de VHippopotame, p. 
246, the raphS-totu-hyoidien, 

In the Flying Squirrel Macalister found that the digastric had two separate 
bellies and a central ronnd tendon which was continued across, from side to 
side above the hyoid bone, as an arch from which the anterior bellies arose. 
This tendinous arch was the sub-hyoidean septum. I have met with a similar 
arrangement in the body of a man, in the University dissecting-room, this 
winter. 

* Gratiolet (p. 299) describes it as 2€ second faUceau of the digastric. It is 
in the plane of tiie digastric ; its mandibular insertion is blended with the inser- 
tion of the digastric, and it occupies the place of the anterior belly of that 
muscle in Man. A hyo-mental is present in some Bats, according to Macalister. 



326 PROF£SSOB HUMPHRY. 

level with the mylohyoid and the genio-hyoid and, partly, is con- 
tinued on into the lateral edge of the hyo-mental. The digcutfic 
(ZX) arises, by a tendon, from near the extremity of the same process 
as the stylo-hyoid, passes internal to the stylo-hyoid, expands into a 
belly which, about on a level with the hyoid, is interrupted by a 
transverse inscription*. In front of this (Z>) it runs parallel, and on 
the same level, with the hyo-mental and is inserted, behind the hyo- 
mental, and continuously with it, into the lower edge of the body of 
the jaw, midway between the symphysis and the angle. 

The facial artery {F, A,) passes, as usual, beneath the stylo-hyoid 
and digastric, and runs along the outer and posterior side of the digas- 
tric, over the side of the jaw, between the digastric and the fibres of 
the masseter which are curling round the anterior border of the pro- 
tuberance to be inserted into its inner side and edge. 

The points shewn by this dissection are, first, the connection of the 
sithciUcmeus colli with, and its termination at, the hyoidean septum. 
Secondly, the presence of a hyo-mental muscle superficial to the 
mylo-hyoid, extending from the hyoidean septum to the symphysis 
of the jaw, and the continuity of some of the fibres of the 9tylo-glo88U8 
with it. Thirdly, the position of the anterior belly of the digastric 
{D\ in front of the inscription, on a level with, and on the 
side of, a part of the same stratum with the hyo-mental, and, 
indeed, continuous with that muscle, and forming a lateral portion 
of it. It is clear that the inscription in the digastric is a part of the 
sub-hyoidean septum, and that the hinder, deeper, belly (ly) of the 
digastric is, by the medium of this inscription, continued into the 
lateral part of the hyo-mental, which forms the anterior belly; just 
as some of the fibres of the stylo-hyoid are continued into the middle 
part of the same muscle. In Man the lateral portion of this hyo- 
mental is absent; and it is the median portion which forms the 
anterior belly of the digastric. Hence the insertion of the digastric is 
near the symphysis of the jaw. The shifting insertion of the digastric 
into the jaw is thus explained by the circumstance that its anterior 
belly may be formed by different parts of the hyo-mental in dijfferent 
animals; and the inscription, usually observable, even when the 
muscle is straight, indicates the line of connection of the portion of 
the deep, or masto- hyoid, and the superficial, or hyo-mental strata 
which combine to make up the muscle. The hyo-mental portion, 
anterior to the inscription, or remnant of the sub-hyoidean septum, 
is a part of the superficial brachio-cephalic stratum ; and the maajx)- 
hyoid portion, behind the inscription, is a derivative from the deep 
brachio-cephalic stratum. 

^ Gratioletf whose aoootmt oorresponds in other respects with that in the 
text, obBerres that the masonlar fibres are not intermpted by any tendinous in- 
scription. It is, howeyer, qnite distinct in the young (8 days old) animal from, 
which my description is taken. 

In the Cat (Fig. 47), as in many others, the digastric is a straight muscle, 
slightly constricted and crossed by a distinct inscription where it passes the 
hyoid, with which it is unconnected. Then it expands to be inserted into the 
whole of the inner sniface of the body of the mandible between the symphysiB 
and the masseter. There is no hyo-mental. 



I 



MUSCLES IN VERTEBRATE ANIMALS. 327 

It is worth while to remark that the hyo-mental in the Hippopo- 
tamus is, together with the anterior belly of the digastric, supplied bj 
the mjlo>hjoid branch of the fifth nerve which runs between them 
and the rajlo-hjoid muscle. 

In the limbless Saurians (Pseudopus, P.) the recttis abdomi- 
nisy which forms the median part of the superficial stratum of 
the ventral muscle in the abdomen, is not separated from the 
cervical {stenuhmastoid) part by an intermediate, thoracic, trans- 
versely disposed (pectoralis) part, but is continued directly on 
into it, or with only some interruption of the deeper fibres by 
the clavicle. In Snakes the foremost bundles of the external 
oblique muscle arise from the mastoid process, and nin back- 
wards towards the ventral scutes beneath which they form a 
superficial rectus. Thus they represent the sterno-mastoid 
muscle in the same manner as do the corresponding fibres in 
Pseudopus P. For a short distance near the head the obliquns 
is overlaid by a superficial, thin muscular sheet extending from 
the cervical spines to the mandible and the scutes immediately 
behind it. This is described as being divided into cermet sub^ 
maadllaris and depressor mandibulce ; and it corresponds evi- 
dently with part of the cervicalis superjicialis of Lepidosiren and 
Dog-fish. But it is not interrupted by branchial openings ; and it 
is not segmented from the subjacent layers of the ventral muscle, 
at least it does not present a stratification from them; although 
such separation may be indicated by the presence of the hyoid 
in the thickness of the sheet (see Fig. 44 aiid description*). 

Traced backwards the superficial (external oblique) stratum 
of the ventral muscle is continued upon the hind limb, with 
more or less interruption by the limb-girdle ; and it ensheaths 
the femoral segment much in the same manner as the humeral 
segment of the fore limb is ensheathed by a projection of the 
anterior prolongation of the stratum'. Ventrally, it is continued 

^ Some of the fibres of this — the cervici suhmaxillaris — ^port of the stratum 
are described by Prof. E. d' Alton (Beschriebung des Muskelsystems eines 
Python biTittatus, Muller's Arehiv, 1884, s. 855) as attaching themselves to the 
hyoid, forming a 'Nackenzongenbeinmnskel.' He also describes a *Biick- 
wartszieher' of the hyoid nmning obliquely and saperfioiaUy from the cervical 
spine to the hyoid. Some fibres of the cervici- or masto-submaziUariB attaching 
themselves to the qnadrate, he calls ' Zoriickzieher' of that bone. It is an 
extension of these upon the mandible which forms the depressor mandibuUs. 

' In certain Snakes the ventral muscles form a funnd-shaped process in- 
vesting the base of the claw, which is the counterpart of the funnel-Bhaped 



328 PBOFESSOB HUMPHRY. 

into the gracilis, which is, manifestly, a serial repetition of the 
pectoralis, and which, like that muscle, is chiefly attached to the 
pre-axial edge of the limb. Dorsally, it is continued into the 
gluteus maximus, which is the serial repetition of the loHssimus 
dorsi, and which, like that muscle, is attached to the post-axial 
edge of the limb and blends with the dorsal muscle (the qiutd' 
riceps) on the first segment of the limb. Both these muscles 
{gracilis and gluteus) incline to the plantar surface of the middle 
and distal segments of the limb, as do the pectoralis and latis- 
sirtms dorsi in the fore limb, and for the same reason. The 
middle part of the external oblique layer is contiuued into 
the tensor vagiruB femoris and the sartoriiuf, which are the 
representatives of the deltoid muscle, and which are cut off, 
wholly or in part, from the obL exL by the ala of the ilium, as 
the deltoid is wholly or in part cut off from the trapezius by the 
spine of the scapula. The opposed edges of the sartorius and 
the gracilis^ are not commonly so approximated to each other 
as are those of the deltoid and pectoraiis- In the interval 
between them, as well as upon them, the oblique muscle is con- 
tinued into the fascia of the thigh ; and the thickening called 
'Poupart's ligament' at the crural arch, where the oblique 
passes into the femoral fascia, is probably a remnant of the 
pelvic septum ; and an ossification in it would form the serial 
homologue of the clavicle*. 

These muscles and fasciae are sometimes covered by an 
extension of the brachio-lateral muscle (p. 320), which may 
{Pteropus) acquire an attachment to the femur; just as in 

myestment carried npon the fore limb of higher ftnimalB and segmented into 
pectortUitt latiaimus dorsi, and trapezio-deltoid. Commonly the interruption of 
the components of the stratum by the pelvic girdle is greater than that by the 
shoulder-girdle. Thus the gracilis is often quite separated from the rest of the 
ventral muscle ; though in some animals, as Cryptobranch, its continuity with 
the caudal and abdondnal parts of the stratum is sufficiently clear : whereas the 
pectoralis is usuaUy to some extent, and often very extensively, continuous with 
the external oblique, or the rectus, or both. So, the sartorius and tensor vagina 
femoris are rarely (they are in Ai) continuous with the external oblique ; though 
the deltoid often is continuous with the trapezius, 

^ In the Babbit the sartorius, arising from Poupart*B ligament, is con- 
tinuous with the gracilis, — ^In Ai, Manis, Pig, and others, the sartorius and 
tensor v, /. are continuous with the gluteus; and in Pig tiie biceps also is con- 
tinuous with them. 

* Probably the marsupial bone, formed from cartilage in the deeper part of 
this septum, corresponds with the sternal end of the clavicle, which, its epiphy- 
sial naoleoB at any rate, is formed in cartilage. 



MUSCLES IK VERTEBRATE ANDCALS. 329 

I* the fore limb the subcutaneous muscle dips down to and is 

I united to the humerus. 

Behind the limb the external stratum is continued upon the 
tail and there blends with the middle stratum. Its superficial 
or subcutaneous layer forms the external sphincter ani. In 
some animals (Hedgehog) it is firmly inserted into the caudal 
vertebrae. 

The followiiig is a review of the serial or sucoessional homological 
relations of the parts of the external stratum of the ventral muscle 
thus far discussed. The gluteus max, is serially homologous with the 
kUiagimus dorsL The tensor vag.f,^ the sariorius^ the femoral fascia 
and part of the ohl. ext,, with the ddtoid and trapezius; Pou part's 
ligament representing the clavicle, and the ala of the ilium represent- 
ing the spine of the scapula. The gracilis with the pectoralis major. 
The middle part of the external oblique and the superficial fibres of the 
rectus with the stemo-cleido-mastoidy and (between the stemo-mas- 
toids) with the mylo-hyoid, the anterior belly of the digastric, the 
hyo-mental and the supeificial fibres of the stemo-hyoid. The rhom- 
boids and serrati postici have no homological representatives; and 
there are no structiures in the tibial and fibular borders of the hind 
limb sufficiently segmented to compare with the tensor plicce alaria 
and the costo-ilaais\ though the sartorius or some of its fibres nearly 
corresponds with the former, and the caudo-pedal with the latter. 
The external sphincter ani may be said to be serially homologous with 
the orbicularis oris; but the subcutaneous muscles generally do not 
admit of homological comparison. 

The relations of the ventral muscles to the alimentary tube are 
therefore as follows : — The internal stratum is gathered round, con- 
tinued into and reflected upon it as levator and retractor ani and is 
gathered round it as post- and sometimes as pre- cardiac diaphragm. 
The middle stratum forms the internal sphincter ani, and is continued 
into the tongue as genio-hyo-glossus and hyo-ghssus, and is continued 
upon the pharynx as thyro- and hyo-pharyngei or constridores. The 
external stratum forms, by its supei^cial layer, the orbicularis oris 
and other circum-oral muscles, and the sphincter ani externum. 

The disposition of the ventral muscle in these three, oblique and 
transverse, planes is a visceral feature, is co-extensive, that is, with 
the visceral region or nearly so. It extends from the anus to the 
head, but not behind the anus, even in Bdellostoma, Snakes and other 
limbless animals. , When the limb-giixUes are present, either with or 
without the limbs, the planes are broken up by and partially lost on 
them ; and when they are not present the planes are blended in the 
caudal muscles. In the Porpoise the recti, and with them the obliqui 
.. intemi and the transversi, diverge and pass, on the sides of the bones 

f supposed to be rudimentary pelvic bones, to the transverse vertebral 

processes in which they terminate about on a level with the vent; 
while the obi. extemi terminate in a defined edge a little in fix)Dt of 



r 



330 PROFESSOR HUMPHRY. 

the bones just mentioned. It will be understood that the stratifica- 
tion is no necessary accompaniment of the visceral region. It does 
not usuaUj take place there in Fishes; and in higher animald it is 
often obscure or imperfect in certain parts, rendering it difficult 
or impossible to decide with certainty from which of the strata a given 
muscle is derived 



Muscles of the Limbs. 

It has already been shewn that limb-girdles are, like the ribs 
and costal cartilages, formed in the transverse intermuscular 
septa of the ventral muscle, and in that part of the thickness of 
the septa which is in the plane of the middle or internal 
oblique stratum. They sometimes gi*ow through the outer 
stratum and project subcutaneously ; and the pelvic bone in 
Snakes lies, beneath the internal stratum, in juxta-position with 
the fascia transversalis. Essentially, however, they belong to the 
middle stratum ; and the muscles passing from them upon the 
limbs may be regarded as derivatives from this stratum, as 
serially homologous, that is, with the muscles passing from 
septum to septum, or from costa to costa, in front and behind 
them. 

It has also been shewn that the external stratum — ^the 
stratum of the external oblique muscle — ^is prolonged upon the 
limbs in the form of a more or less complete funnel-like invest- 
ment of each limb. This is seen in its simplest condition in the 
Snakes that are possessed of claws. It is also well seen in Lepi- 
dosiren, where it consists of a simple sheath, interrupted, it is 
true, in front, by the branchial opening, yet surrounded by and 
extending along the fin ; and different forms of it are exhibited 
in different animals. There are, therefore, derivatives of the two 
outer strata of the ventral muscle, at any rate, contributing, and 
largely, to the muscular basis of the limbs. 

A limb is usually composed of a series of cartilaginous or 
osseous pieces serially arranged and moveable upon one another, 
of which the fin of Lepidosiren is one of the simplest examples ; 
and the instance of the paddle of Ceratodus has shewn that the 
muscular fibres pass from piece to piece, indeed that the muscu- <. 

lar tissue of the limb is, in the primitive form, segmented into 
transverse planes corresponding with the axial cartilaginous, or 



V 



MUSCLES IN yEKT£BEA.T£ AMIHALS. 331 

osseous, segments, thus resembling the disposition of the muscles 
in the trunk, and more particularly in the tail, of simply con- 
structed animals. But, as we have found in the trunk, the 
muscular fibres, the superficial fibres more particularly, are often 
not confined to their particular segments. They commonly 
break through the intervening barriers, or intermuscular septa, 
and range on to more difltant segments, blending -with the 
muscles of those segments. Hence the muscles of any division 
of a limb consist, usually, of three layers. First, and deepest, 
are the fibres of the segment itself, the 'intrinsic* fibres ; of these 
the proximal series are the bundles passing from the girdle to 
the first segment of the limb : secondly, the fibres derived from 
distal segments, the 'extrinsic' fibres; and thirdly, and most 
superficially, the fibres derived from the ventral muscle, the 
superficial 'ventro-appendicular' fibres. The components of these 
three layers are blended together in a variety of ways, render- 
ing it often difiicult or impossible to distinguish to which layer 
they appertain. 

In the simplest condition, as that of Lepidosiren, the carti- 
laginous or osseous pieces of the limb are joined to one another 
by simple tissue, and are moveable in any direction ; and the 
muscular fibres are uniformly disposed around the joints, the 
deepest, or ' intriDsic', fibres forming a sort of circular capsule, 
and being blended on the exterior with the 'extrinsic' and the 
' ventro-appendicular' fibres. When movements in particular 
directions are required, the conformation and structure of the 
joint is proportionately modified, and the muscular fibres are 
arranged and segmented, more or less distinctly, into bundles 
or muscles to effect the object. In the case of the proximal 
joint of the limb — ^that of the first cartilage or bone with the 
girdle — ^the movement is, on the whole, circumductory or in 
any direction ; and the muscles are disposed around the joint in 
a nearly circular manner. Usually, however, even here, certain 
movements take place more frequently or more freely than 
others ; and the muscular fibres are accordingly arranged in 
groups, or muscles, having more or less independent action. In 
the distal parts of the limb the movements are commonly much 
more restricted, are limited indeed, nearly or quite, to one 
plane, which is, speaking generally, the same for all the joints 



332 PROFESSOR HUMPHRY. 

of the limb. They all admit cf flexion towards the palmar 
or plantar or ventral aspect, and of extension towards the 
dorsal aspect. Supposing the limbs stretched, as in their 
primitive condition, horizontally in straight lines and at right 
angles, from the trunk, then all the joints admit of flexion 
towards the ventral surface of the trunk and of extension to the 
horizontal line. At the wrist and ankle the extensor move- 
ment may, in some animals, be carried beyond that line (in 
Man the foot has come to be placed at a right angle with the 
leg) ; and the same is the case at the joints with the girdles 
where, as just said, other movements are also permitted. In 
all the joints, however, except that with the girdle, the movement 
is nearly restricted to the plane indicated. Hence the muscles 
are arranged in two sets, one upon the ' flexor' or 'ventral' aspect, 
which may also be called 'palmar' in the fore limb and 'plan- 
tar' in the hind limb, and the other set upon the 'extensor' 
or 'dorsal' aspect; and there is, speaking generally, a corre- 
spending arrangement of the nerves. 

It must not, however, be supposed that the arrangement 
is rigidly adhered to. Marginal portions of the ventral series 
occasionally (witness the lunibricales) stay upon the dorsal 
aspect and serve as extensors ; and more frequently marginal 
portions of the dorsal series incline over the sides and upon the 
ventral aspect and serve as flexors, and perhaps become united 
with the flexors. 

That there is a similarity, a general homological correspond- 
ence, between the muscles of the fore and of the hind limbs, as 
well as between the limbs of different animals, is self-evident. 
Their outer sheets are projections of serially homologous portions 
of the ventral muscle carried upon similar outgrowths from serially 
homologous ossifications in the ventral intermuscular septa ^. 
The dorsal muscles of one limb and in one animal correspond, 
though not unexceptionably, with the dorsal muscles of the 

^ The sepia in wMoh the oorresponding limb-girdlefl are fonned are not, 
nmnerioally, the same in the different orders of vertehrates. The soapnlar 
girdle, for instance, in the Fish is developed in a septum dose to the head ; 
whereas in the Bird it occupies a septum far removed from the head. So, the 
position of the pelyic girdle, though presenting a steady relation to the hinder 
part of the visceral cavity and the termination of the alimentary canal, varies 
greatly in its distance from the scapular girdle, ovdng to the variation in the 
number of intervening muscular and intermuscnlar pluies in different animals. 



MUSCLES IN VERTEBRATE ANIMALS. 333 

other limb and in other animals ; and the same with the ven^ 
tral series. There is, moreover, often an antagonistic similarity 
between the ventral and dorsal muscles, according with the 
similarity in the action requisite to produce the flexor and the 
extensor movements. In short, the similarity of muscular 
disposition about the several joints, in the same or the different 
limbs of the same or different animals, is, as we might expect, 
generally proportionate to the similarity of the form, move- 
ments and position of the joints ajid of the force required \ 
Differences in these. are sometimes, indeed not unfrequently, 
associated with such varieties in the combination of embry- 
onic germs as to defy attempts at an exact homological com- 
parison. 

Two important features of difference which serve to modify 
the muscular relations in the two limbs at once suggest them- 
selves as being present to a greater or less degree in most 
animals. First, the superficial position and the mobility of the 
shoulder-girdle, as compared with the pelvis; and associated 
with these are the variability and instability of the subgl^ioid 
parts of this girdle, and the frequent presence of a clavicle. 
The shoulder-girdle and its musdes often, indeed commonly, 
deviate from their simple primitive form and relations much 
more than do the pelvic-girdle and its muscles ; and they are 
liable to much greater variations in position — ^fore and aft and 
in the depth of plane at which they are situated — and in their 
relation to the other parts of the osseous skeleton. In one 
animal the shoulder-girdle is joined to the head, giving it^ in 
the opinion of some anatomists, a claim to be regarded as an 
appendage to the skull; in another animal it is joined to the 
vertebral spines ; in a third to the vertebral bodies ; in a fourth 
to the sternum ; and in a fifth it is free from all these. These 
varieties in the girdle involve a greater amount of variety in 
the muscles connecting it with the rest of the trunk and passing 
from it to the limb^ than is the case with the muscles of the 
pelvis. There is, however, in connection with the latter^ the 
varying development of the tail, which leads to considerable 

^ I do not mean to aBsert that all mnsoular Yarieties are thna teleologicaUy 
determined; bat it is very generally so; and it is not easy to adduce onques- 
tionable exceptions. 

VOL. VI. 23 



834 PEOFESSOR HUHFHBT. 

variation in the muscles passing between this part of the body 
and the pelvis and hind limb. 

The second great feature of difference is the difference in 
the rotation of the two limbs ; the dorsal surface of the thigh 
and knee being directed forwards, while that of the arm and 
elbow is directed backwards \ In the hind limb the leg and 
foot are involved in the same rotation as the thigh and knee. 
But the forearm and hand undergo a rotation in a different 
direction to that of the arm by means of the proation of the 
radius ; so that the dorsal surface of the hand is directed the 
same way as the dorsal surface of the foot. The effect of these 
rotations is that the radial — or pre-axial — edge of the humerus 
looks outwards, and the tibial — or pre-axial — edge of the femur 
looks inwards. This leads to a difference in the insertion of 
some of the otherwise homologous muscles at the upper ends 
of the two bones, for instance, the radial tubercle of the hume- 
rus presenting on the outer side of the limb, receives the mus- 
cles from the dorsal aspect of the scapula and exceeds the idnar 
tubercle in size ; whereas the fibular tubercle of the femur 
rises into strong relief beneath the dorsum of the ilium, and 
receives the muscles which arise from that surface and exceeds 
the tibial tubercle in size. 

I may obserye that such a variation in the locality of the inser- 
tion of correspondiDg tendons in the two limbs, and indeed in the same 
limb in different animals, is by no means uncommon. It is seen in 
the instances of the extensor muscles of the leg and foreanfk, in the 
exteiuorea carpi radiales, the biceps brachiif IcUissimtis dorsi, &c. Too 
much importance has, I think, heen attached to the insertion of a 
muscle as a guide to its homology : and the statement that the inser- 
tion is more to be depended upon, in this respect, than the origin is 
scarcely supported by observation. 

We have already (p. 327) traced the ventro-appendicular 
muscles from the ventral muscle to the limbs, and pointed out 
their serial homological relations. It remains to follow them 
upon the limbs. Normally, or primitively, they spread, as in 
Lepidosiren, and less distinctly in Ceratodus, over the whole 
surface of each limb, forming an outermost muscular cover- 
ing of the limb; and they are still seen to expand upon the dis- 
tal segments in many even of the higher animals, especially in 

^ See my Ob$, on the Limbs of Vertebrate Animdl$t p. 16. 



V 



MUSCLES IN YEBTEBBATB AKDIALa 835 

the hind limb. But they are often reduced, in their course down 
the limb, to fasciad or areolar fibres, and often are arrested by 
insertion into a projecting process of bone at some higher point; 
or they blend with, and are lost in, the subjacent strata. 

To take first the 'palmar' and 'plantar', that is, the ^ven- 
tral' divisions of the ventro^appendicular muscle, which we call, 
respectively, pectorcdis and ffracUis, Each extends along the 
pre^axial edge of its limb, inclining to the palmar, or the plan- 
tar, surface. Each reaches sometimes to the distal segment, 
but is usually interrupted or lost eai*lier; and each blends in 
variable degrees with the subjacent muscles. The pectorcdis is 
commonly arrested at the radial tubercle of the humerus; but, 
sometimes (Orycteropus), part of it accompanies the bicq[>8 to 
the radius, or (Otter and Wild Cat), together with some fibres 
of the trapezio-deltoid, accompanies the brachiaiia cmUcua to 
the ulna, or (Seal) expands into the fascia of the forearm and 
so reaches the hand. It is, in some instances, free firom the 
subjacent muscles, and in others is blended with the coraco- 
brachials, or is connected with them by means of the pectoralia 
minor (see p. 344). The gracilis is rarely, if ever, arrested at so 
early a point as the pectoral. It is commonly inserted into the 
tibia, occasionally (Pteropus) blending with the semitendinosTis, 
and, now and then (Unau and Al), joining the biceps flexor cruris 
and acquiring an attachment to the fibula. It often extends 
down the inner side of the tibia, over the ankle and foot, spread- 
ing upon the dorsal and plantar surfaces of the foot (Seal). It 
is in some instances free from the subjacent adductors; in 
others (Cryptobranch) it is scarcely segmented from them. 

The 'dorsal' divisions of the ventro-appendicular muscle 
have often a wider range than the ' ventral ' both in their con- 
nection with the trunk and in the area of the limbs which 
they cover; and they are commonly sectorially segmented in 
some degree. The dorsal surface, as seen most distinctly in the 
fore limb of Dog-fish (Figs. 28 and 30) and in the hind limb of 
Ceratodus (Fig. 34), and the post-axial edge of the limb, may 
be regarded as their legitimate destination; but they not unfre- 
quently spread, over both the pre-axial and the post-axial edges, 
upon the ventral surface of the limbs. They are, consequently, 
attached to various points. In the cylindrical (Lepidosiren), or 

23—2 



836 PROFESSOR HUMPHRY. 

rudimentaiy (Snakes), state of the limbs, they are not seg- 
mented from the pabnar and plantar portions of the sheet, and 
are not themselves longitudinally or sectorially cleft. 

In the fore limb there are usually two sectors of the dorsal 
division; one — ^the trapezius— converging from the dorsal aspect 
of the head and of the front of the trunk, and the other — ^the 2a- 
tissimris dorsi — converging from behind, upon the limb. The tra- 
peziys sector in branchiate animals (pp. 259, 273) is interrupted 
and devoted to the gill apparatus, and scarcely reaches the 
limb. In some animals, devoid of gills (Cryptobranch), it does 
not extend beyond the anterior edge of the scapular and the cora- 
coid parts of the girdle. In others (certain Mammals) it runs 
on to the radial tubercle of the humerus, or to the radius, or 
even (Otter^) to the ulna, joining the flexors of the forearm. It 
usually presents an inscription as it passes over the shoulder. 
This inscription is the superficial part of the septum in which 
the girdle is formed; and, in Mammals, the anterior edge of 
the scapular part of the girdle, called the ' spine \ commonly 
grows through the septum, so occupying the inscription, and 
bisecting the upper or supra-scapular part of the muscle, trans- 
versely, into posterior, or deltoid, and anterior, or trapezius 
parts. In the lower, or coracoidal, part of the septum the case 
is somewhat different. Sometimes the inscription {i,e. the 
remnant of the septum in the muscle) remains, and marks the 
division into trapeziuB and deltoid parts. Sometimes it is 
nearly obliterated. Sometimes it becomes ossified into a ' dar 
vicle' or 'epi-coracoid' (p. 316), the ossification being (many 
Carnivora) confined to the deeper tract of it and limited in 
extent, or extending through the thickness of the muscle, and 
ranging, from the acromial end of the spine of the scapula, to 
the sternum*. The most important varieties, therefore, in this 

> In an Otter the foremost fibres of the trapezius, oontinned as deltoid with* 
an interrening inscription, and aooompanied oy some fibres of the pectoralit, 
descended in front of the bracfUalU anttcus to the nhia. 

* In Lizards the daTide extends along the anterior edge of the scapula to 
its base. In Ai it is attached to the ooraooid. In Fidies it is large, closely 
applied upon the ooracoid, and meets its fellow in the middle line ; but it is not 
eertain that ossification ever extends from the ooracoid into it. It seems in 
this respect to bear to the coraooid somewhat similar relations to those which 
the supra-, or better epi-, scapular bones (Parker's supra-davioles) bear to the 
scapula. I haye spoken of the clavicle as an ossification in one of the muscular 
septa of the brachio-cephalic stratum, corresponding wiUi the epicostals. But 
it should be added that it is situated at a deeper level than they, in the deeper 



MUSCLES IN YEBTEBRATE ANIMALS. 837 

trapezio-deltoid sector are; firsty that in which it does not ex* 
tend below the anterior edge of the girdle; eecoTicUj/f that in 
which it extends beyond the edge of the girdle and its septum 
down to the humerus or forearm; thirdly, that in which it is 
divided transversely, in part or the whole of its width, into 
two distinct muscles by ossification extending from the edge of 
the scapular part of the girdle into the septum which traverses 
it and by ossification arising independently over the coracoidal 
part of the girdle. The portion of the trapezius which, in 
Cryptobranch, is inserted into the anterior edge of the scapula 
is, in Mammals, continued, beyond that edge, which grows up 
as the spine, into the scapular portion of the deltoid; and the 
portion which, in Cryptobranch, is inserted into the coracoid is, 
in Mammals, continued, beyond the coracoidal septum, or the 
clavicle which may form in it, on into the clavicular portion 
of the deltoid. These two portions of the deltoid are not unfire- 
quently separate. 

The sector in the hind limb, corresponding to the trapezio* 
deltoid, is recognisable as a distinct element only behind the 
pelvic girdle, where it forms the aartorius and the tensor vagince 
femorie. These are usually cut off from the extemaX oblique 
part of the ventral muscle lying in front of them, which, or 
part of it, may be supposed to correspond to the trapezius, by 
the edge of the ilium; and they thus answer, serially, to the 
scapular portion of the deltoid. The sartorius is sometimes 
(Unau and Ai), like the deltoid, inserted into the pre-axial 
edge of the first bone of the limb; but, usually, it runs into the 
second bone. In other ^ords, the sartorius usually continues 
its course on to the tibia, but is occasionally arrested at the 
femur*; whereas the deltoid, or part of it, occasionally (Orycte^ 

instead of in the superficial part of the Btratmn, as seen in Ps. Pallasii, and 
also in Camiyora, where the rudimentary olaTiole is found in the deeper part of 
the septum between the trapezius and the deltoid. It has not unfrequently 
conneetions with the muscles of the subjacent stratum — the tubclavius and the 
stemo- and omo-hyoids. It is extended between two cartilage bones, of which 
one (the sternum) is in a deep level. According to Gegenbaur, it is not without 
claim to a cartilaginous origin itself at one part; and it has a cartilaginous 
epiphysis. In short, it appears to afford an example of remarkable blending of 
the superficial membranous and the deeper cartilaginous ossifications, the 
former preponderating from a very early period, and in some instances, probably, 
constituting the only basis of the bone. 

^ The sartoriut sometimes (Hippopotamus) runs down the middle of the 
dorsal aspect of the thigh, over the patella, and is lost there. In the Heron it 



S88 PROFESSOR HUMPHRY. 

njpoB) continues its course on to the radius, but usually is ar- 
rested at the humerus. The coiurse of the tensor v, /. into the 
fascia does not distinctly correspond with anything in the fore 
limb. It is present only in Mammals, and is often wanting in 
them. The aariorius may (Rabbit) extend, along Poupart's 
ligament, to the gracUta, as the deltoid often extends, aloDg the 
clavicle, to meet the pectoral. This is, however, rare; and the 
intermediate space between the two muscles in the retiring 
angle of the bend of the groin, where there is no ' point d'ap- 
pui' for muscular action, is usually occupied by fascia. That 
fascia is^ accordingly, the serial representative of the clavicular 
portions of the pectoral, deltoid, and trapezius, Poupart's liga- 
ment being the serial septal representative of the clavicle. 

The posterior sector of the dorsal portion of the ventro- 
appendicular muscle in the fore limb — the latissimus dorsi — is, 
like the pectoral, a very general appurtenance to the fore limb 
in animals above Fishes. Even in Fishes it is sometimes (Dog- 
fish, Fig. 28) represented by a thin stratum segmented from 
the ventral muscle and passing upon the dorsum of the fin, the 
pectoral being in like manner segmented and passing upon the 
palmar surface of the fin. In Lepidosiren (Fig. 24) we have 
seen the latissimus dorsi and the pectoralis travelling forwards 
as a continuous sheet constituting the hinder part of the super- 
ficial brachio-cephalic stratum. In this animal the laUssimus 
dorsi is attached to the scapular part of the girdle in addition 
to passing upon the fin ; and in Dog-fish it is barely segmented 
from the deeper {serratus) part of the stratum which is also at- 
tached to the scapula. In Man, and some other animals, it still 
retains the scapular connection; and, as before (p. 319) re- 
marked, it often retains its connection with the pectoral by 
axillary bundles passing from it to that muscle. It is traceable 
to variable points in the limb. In Lepidosiren its fibres run on 
with others to the extremity of the fin. In TJrodelans it is 
blended with the triceps; and more or less of this union is often 
found in Mammals. In Saurians and Birds it passes, between 
the scapular and humeral origins of the triceps, to the radial 
edge of the humerus. In Mammals, passing internal to the tri- 

Ib more sepaiate from the glutew than in moBt Birds, and some of its fibres are 
traoeable into the gaetroenemiut. 



MUSCLES IN VSRTEBRATE ANIMALS. 839 

oepsy it is inserted into the ulnar edge of the humerus, sendiiiig 
often a dorsi-epitrachlien down to the uhiar condyle, or (Cyclo- 
thurus) extending along the fascia of the forearm to the ulnar 
side of the carpus ^ or (Manis) running into the flexor sublimia 
digitorum, or (Rabbit) into the flexor carpi tdnarie. 

This muscle thus affords a very interesting illustration of the 
varying points at which a ventro-appendicular muscle becomes 
arrested in different animals, and the varying muscles of the deeper 
strata with which it becomes blended or, rather, with which it retaing 
its connection. 

It is worthy of remark that the fibres of this muscle, and also of 
the pectoral and, though less frequently, those of the deltoid, often cross 
one another in a remarkable manner; those which arise most pos* 
teriorly passing behind the others to an insertion in fit^nt of them, bo 
cansing a sort of twisting or folding-in of the lower edge and, some- 
times, leading to a cleavage in the plane of the muscle. I have 
before called attention to this point (p. 296aiiidJoum(UofAn(U, TV. 35). 

The corresponding sector in the hind limb — ^the ffluteui 
mctxinrns — ^has relations as varying as those of the tatiesimus 
dorst. Its connection with the ventral muscle is well exempli* 
fied in Dog-fish (Fig. 30), and Ceratodus (Fig. 34), as well as the 
mode in which, in a primitive condition, it extends upon the 
limb, and blends with the proper muscles of the limb*. From 
these animals upwards the connection with the ventral muscle 
rarely reappears*. Thenceforwards its chief attachment is to 
the ilium (which appears through the ventral muscle like the 
spine of the scapula) and to the neural spines and arches of the 
caudal vertebrae. In the lower animals it is often, as we have 
seen in Cryptobranch, so united with the extensor cruris as to be 
with difficulty distinguished from it; and in Birds it blends 
with that muscle in the thigh. The same connection exists, 
to a greater or less extent, in Mammals; though sometimes, aa 

^ Bepresenting the eoito-aUirU of the Bixd (p. 819). 

* It is not quite correct to speak of the mnsole referred to In these animals 
as the glvXew sector only; forasmaoh as it represents, rather, the entire on* 
divided dorsal portion of the yentro-appendicnlar sheet. In Mammalfl also it is 
often nndiyided. 

In anonrons Batraohians and Sanrians, this dorsal portion is a not dis- 
tinctly segmented mnsde. 

' In Manis the dorsal portion of the yentro-appendicnlar mnsde is largely 
developed, blending above with the external obliqae and reaching to the lumbar 
spines, and below ertftnding over the buttook and the fore and outer parts of 
the knee and thigh. 



340 PB0F£SSOB HUHPHRT. 

in Man, the connection is with the fascia covering the qiiadri- 
eeps. like the kttissimus d. it is, in addition, often inserted 
into the post-axial line of the femur ; but it does not follow its 
homologue by occasionally passing, between the divisions of 
the extensor muscle, to the pre-axial line. It sometimes ranges 
along the margin of the ilium to unite with the tensor v. /. and 
the sartorins, so obliterating the division between the two 
sectors; and on the limb it often reaches the fibular malleolus 
and side of the foot. 

Before concluding the account of the ventro-appendicular 
muscles, it is necessary to mention certain derivatives from the 
ventral muscle of the tail which belong to this series, some of 
which are not represented in the fore Umb. 

Of these, the first and most superficial is that which I 
named Caudo-pedal in the description of the Cryptobranch« 
It occupies an intermediate position between the gluteus and 
the gracilis, and descends along the plantar surface of the 
Umb to the distal parts, blending with the flexors of the digits. 
In Fishes it is not distinctly represented ; and in animals above 
Reptiles it is lost or represented only by fascial tissue in its 
upper part. Its lower part in these animals is probably repre- 
sented by the plantar fascia, or the superficial layers of the 
plantar fascia, and, perhaps, by the superficial layers of the 
tendO'AchiUis and by some fibres of the gastrocnemius. In the 
fore limb of Birds it seems to be serially represented by the 
costo-alaris, or it$ superficial fibres, which, as I have before 
said, is the antagonistic homologue of the cervico-alaris {tensor 
pliccB alaris). Such antagonistic homologue to the caudo- 
pedal in the hind limb we do not find. It would be represented 
by fibres from the external oblique, between the sartorius and 
the gradUs, passing down to the distal region of the limb\ 

Beneath, and connected with, the caudo-pedal is the caudo- 
arural of Cryptobranch, which fuses with the adducto-flexor 
mass passing to the leg, and more particularly with that part 

1 In Crocodiles a large muscle passes from the hindmost projecting point of 
the ilium; and at the ham its tendon is, partly, connected with the deeper 
muscles and, partly, extends superficially down to the foot. A segment of it 
internally joins the gradlU. ExtemaUy, it is in contact with the bieep9 and gUi- 
teo-rectui, I suppose it is the representative of the caudo-pedal, starting from 
the ilium as a fixed basis instead of from the caudal vertebne. 



MUSCLES IN VERTEBRATE ANIMAIS. 841 

of it which represents the semitendinosus, thus giving rise to» 
or causing the persistence of, an inscription which I have 
(p. 19) supposed to represent the inscription found in that 
muscle in Man and some Mammals \ The only trace of serial 
homologue in the fore limb to the caudo*crural would be fibres 
of the hinder portion of the pectoral, or fibres of the oblique 
or rectus, joining the biceps brachiL But none of them so 
coalesce with that muscle as to cause an inscription in it. 

Both the caudo-pedal and the caudo-crural lose the trans- 
verse inscriptions where they separate from the ventral muscle 
in the tail ; but there is in the retiring angle between the tail 
and the hind limb in Saurians a thick transverse fascial band, 
which I suppose to be an inscription between the caudal and 
the crural parts of these ventral, or ventro-appendicular, 
muscles, and which seems to correspond, antagonistically, with 
Poupart's ligament, and so, serio-antagonistically, with the 
clavicle. 

Still deeper, and belonging to a deeper plane, is the caudo^ 
femoral, which is usually present in ovipara above Fishes, and 
is sometimes designated agitator caudal. It is inserted into the 
femur, and is, in varying degrees, blended with the deep 
muscles of the thigh — the addAwtora and the aemimevnlbra'nomB 
— and sometimes with the origin of the flexors of the foot and 
toes, affording an interesting example of the prolongation of 
the deeper strata of the ventral muscle to the distal parts of 
the limb. 

The caudal derivations, which form so important an element 
in the muscular system of the hind limb of Urodelans and 
some B.eptiles, are present, though less strongly developed, in 
Birds, and are not uncommonly met with in Mammals. Thus 
the semitendinosvs sometimes derives its chief origin from the 
sacrum. In the Otter and Racoon, the caudo-^yrural is more 
distinctly represented by a muscular slip derived partly from 
the spinous and partly from the transverse processes of the 
caudal vertebrae. It joins the hinder surface of the semitendi' 
nosus, and is prolonged with the posterior fibres of that muscle 

1 In Bome large-tailed Mammals (CyelothamB and Mania) the $emitendino$ui 
retains the caudal ae weU as the pelvic factors. I have not, however, found the 
inscription in any of these. 



342 PROFESSOR HUMPHRY. 

to the inner side of the heel. The cando-femoral is also repre- 
sented in the Otter by a thin muscle^ arising from the trans- 
verse processes of the caudal vertebrsB, passing internal to the 
sciatic nerve^ and into the middle third of the linea aspera, 
between the gluteus and the adductors and distinct from both. 
The pyri/ormis and quadratas femoris muscles are both present 
in this animal In the Rat a caudo-femoral muscle passes from 
the sacrum, beneath the gluteus^ to the internal condyle and the 
post-condyloid bone, between the addr. m, and the semdmembra- 
no8U8. It lies near to, and in the same plane with, the ischio- 
eattdal. The antagonistic homologues of the two {caudo-femoral 
and isckixhcaudal) are probably the psoas magnus with its 
neighbour the psoas parvus. 



The Muscles passing from the Limb-Oirdles to the 

Limbs 

may be regarded aa extensions of the deeper strata of the 
ventral muscle — as deep ventro-appendicular muscles— forafl* 
much as the girdles are ossifications in the deeper level of the ^ 

ventral inter-muscular septa. They share the surface of the 
girdles with the fibres of the ventral muscle which pass back- 
wards or forwards to the adjacent trunk segments. Hence the 
extent of their attachment to the girdles varies inversely with 
the extent of attachment of the direct fibres of the ventral 
muscle. Where the latter are numerous, as on the ilium of 
Cryptobranch, the muscles passing from the bone to the limb 
are few. Where, on the contrary, the range covered by the direct 
ventral fibres is small, as in the pubischium of Cryptobranch, 
the size of the muscles passing to the limb is considerable. In 
like manner the extent of their attachment to the first segment 
of the limb reciprocates with that of the origin of the muscles 
passing to the second segment. Thus, in Cryptobranch, the 
pecUneus engrosses the whole of the dorsal surfeice of the 
femur; whereas in most animals that surface is occupied by 
the extensor of the leg, and the pectineus is compressed within 
narrow limits. It may be inferred that the corresponding 
embryonic feu^tors are, in some instances,, employed in the 



MUSCLES IN VERTEBRATE ANIMALS. S43 

building up of the one series of muscles and, in other in- 
stances, of the other series. 

These muscles surround the joint of the first limb-segment 
with the girdle in a more or less circular, or capsular, manner. 
Usually, however, they are arranged in two divisions, or groups, 
in each limb. The one group passes from the outer, or posterior, 
surface of the girdle, beneath the joint, to the ventral or under 
surface of the limb, constituting a palmar or coraco-humeral 
series in the fore limb and a plantar or pubischio-femoral series 
in the hind limb. The other group passes from the outer, 
or hinder, surface of the girdle, above the joint, to the dorsal 
surface of the limb, constituting a dorsal or scapulo-humeral 
series in the fore Hmb, and a dorsal or iUo-femoral series in the 
hind limb*. These groups do not adhere strictly to their 
respective limits. They sometimes extend from the outer 
surface, over the margins, upon the inner surface of the girdles; 
and they sometimes extend from the area of the girdle beneath 
the joint to the part above it, and vice versd. Moreover, in 
accordance with the difference in the rotation in the two limbs, 
the insertion of corresponding muscles may be on the one — 
the pre-axial or radial — side in one limb, and on the opposite — 
the post-axial or fibular — ^side in the other limb. It not unfre- 
quently is found, in the case of each of the several groups, that 
it is imperfectly segmented from the superficial ventro-appendi- 
cular stratum on the one surface, and from the muscles which 
pass on to the next segment of the limb on the other surface. 

In animals above Fishes the OORACO-HUMERALS, or, cus they 
are more generally called, coraco-brachials, are commonly 
divided into segments which vary in number and size with the 
number and size of the coracoid processes ; and they are some- 
times absent when these processes are abortive, as in Mole, 
Cyclothurus and Seal. They arrange themselves in two di- 
visions. First, those which lie superficially with regard to the 
biceps brachii muscle and which pass to the radial tubercle 

^ These are seen in their simplest form in Geratodus (Fig. 34), where the 
ooraco-brachial and the scapulo- brachial mnscles are seen extending from the 
respeotiye parts of the girdle upon the ventral and dorsal surfaces of the pec- 
toral fin. The arrangements are similar in the hinder fin ; and there are addi- 
tions from the ventral muscle. " 



844 PROF£SSOB HUMPHBT. 

of the humerus immediately above the level of the pectoralis 
major and also extend beneath that muscle. These constitute 
a superficial or pre-axial division. Secondly, those which lie 
beneath the biceps and pass to the ulnar tubercle and ulnar 
side of the humerus. These constitute a deep or post-axial 
division. 

The superficial division comprises the epicoraco-htmteral 
and the precoraco-humeral, and probably the suprorspinatus, 
the levator humeri and the pectoralis minor. All these^ with 
the exception of the last, which I will consider separately, may, 
I think, be regarded as parts of one muscle which occupies, 
essentially, the precoracoid process and might be called the 
precoraco-humeral muscle. It may spread upon the edge of 
the coracoid on the one side, and be segmented, so forming 
an epicoraco-hvmerdl ; and it may spread upon the supra- 
spinal space of the scapula on the other side, so forming a 
supraspinatus. In Cryptobranch (p. 52), where the precora- 
coid processes are laige, the precoraco-humeral muscle is 
well developed; and the epicoraco-humeral, in the same plane 
with and scarcely segmented from it, spreads over the edge 
of the coracoid ; but there is no supraspinatus. In Meno- 
branch the latter muscle is present, in addition to the other 
two; but its edge is continuous with that of the precoraoo^ 
humeral. In the Seine the precoraco-humeral and the supra^ 
spinatv^ are large and separate; but there is no epicoraco- 
humeral extension of them over the broad origin of the biceps^ 
In the Crocodile the precoraco-hum^eral is large, arising from 
the precoracoid process — ^from the outer surface and the anterior 
edge, and also from the deeper surface of that process — ^much as 
the pectineus often arises from the corresponding surfaces of the 
pubes. It is nearly on a level with pectoralis on the one side, 
and the deltoid on the other, and is inserted between them. The 
suprorspinatus is small, lies partly beneath it, and is imperfectly 
segmented from it. There is no distinct epicora^co-hwrneral, 
the suiface of the coracoid, internal to the origin of the biceps, 
being closely covered by the pectoralis which derives some 
fibres from its edge, from the place, that is, where the epicoraxx>' 
humeral usually arises. In the Bird the precoracoid process 
does not run out distinctly. It is bent down with the coracoid 



MUSCLES IN VERTEBRATE ANIMALS. 345 

to the sternum, forming an acute angle with the sternum ; and 
the precordcO'humeral muscle follows it, together with the 
epicordcO'humeral factors. These, or some of them, extend upon 
the sternum and form the levator humeri or pectoralts tertiTis^, 
as it is sometimes called. In Mammals the suprorspindtua and 
the pectoralis minor are the representatives of this division. 

The fibres of the epicoraco-humeral part of this superficial, 
pre-axial, or supra-bicipital, division of the coraco-humerals 
lie immediately beneath the pectoralia m4ijor in its whole course. 
I have remarked that in Crjptobranch its superficial fibres are 
blended with the under surface of the pectoral, and that in 
Crocodile the fibres that correspond with it form part of the 
origin of the pectoral It thus, to some extent, occupies the 
place of the pectoralis minor; and if we suppose it continued 
upon the under surface of the pectoral, and in variable degrees 
segmented from that muscle, it would quite correspond with 
the ordinary mammalian pectoraiis minor, the proper insertion 
of which appears to be the radial ridge or tubercle of the 
humerus. It is, however, in Man and some animals, arrested 
wholly, or (Rat), partially, at the coracoid and is often quite 
segmented from the pectoralis major. Thus, I conceive the 
pectoralia minor to be formed from factora of the pectoralis 
mujor, which, or some of which, represent the epicoraco-htrnie- 
ral of Urodelans, Reptiles and Monotremes, and that it also in 
part represents, indeed is the nearest representative of, the 
levnior humeri of Birds*. 

^ Also the secundus when both are present. 

s This view is in aooordance with the faot that the levator humeri sometimes 
absorbs the iupra-spinatus, or derives an accession of fibres from the npper 
surface of the soaptQa, as weU as with the occasional passage in Man, as noted 
by Macalister, Wood and ottiers^ of the pector<ili$ minor over the coracoid 
process beneath the coraco-acromial ligament where it has been found to blend 
with the supra-tpiiiatuSf or with Ihe capsule of the joint, or to pass directly to 
the head of the numems. It harmonizes also with the close relation of the 
tupra-^natw to the tendon of ^e biceps, a relation corresponding with that of 
the epieoraco-humeral to the bieepe in Urodelans and Reptiles. 

It however is not quite in accordance with the view of Prof. Bolleston, Trant. 
LimL 8oe, xzvi. 609, that the euhclavius is the homologne of the levator 
humeri. I have before (p. 815) given reasons for thinking that the tuhclavius is 
an extension of the middle stratum of the ventral mnsde and is the serial homo- 
logue of the costo-eoracoid muscle. It lies in front of, or above, the costo- 
ooraco-clavioular ligament ; and when it is prolonged beyond the range of the 
eiavide, it usually expands upon the supra-tpinaUu or the dorsum of the 
scapula, forming the stemo- or atemO'CostO'ScamUarii, as it does in Hippo- 
potamus and some other animals. Sometimes, nowever (Pig), it reaches the 
radial tubercle of the humerus ; but its destination is rather to the dorsum of 



346 PROFESSOB HX7MPHBY. 

>The deeper, post-axial, or sub-bicipital, coraco-humerals 
arise from the coracoid beneath the biceps, as best seen in Rep- 
tiles. They may be traced, in these animals, taking origin 
from the under and hinder surfiioe of the coracoid and spread- 
ing upon the under surface of the scapula, where a portion of 
them forms the svhscapyiaris. In Mammals, this muscle 
is quite segmented from the others \ They are inserted into 
the ulnar edge of the humerus; it is inserted into the ulnar 
tubercle. The one next below the srsbscapilar — ^the coraco-6r.- 
wMiua — is generally present and is inserted into the middle of 
the shaft. The passage through it of the external cutaneous 
or musculo-cutaneous nerve indicates a tendency to division ; 
and in several Mammals (Babbit, Proboscis Monkey and Jerboa) 
the upper segment is inserted separately into the ulnar tubercle 
forming a swperwr caraco-brOfChiaL In Amphibians, Reptiles, 
and Monotremes there is commonly a third segment, an inferior 
coraco-brcLohial, which extends to the ulnar condyle; and the 
brachial artery with the median nerve passes between it and 
the middle coraoo-brachial. 

The PUBISOHIOFEMOBALS are less variable in number and 
size than their homologues just described, owing to the greater 
uniformity of the pubic and ischiatic bones in comparison with 
the coracoids. Like the coraco-humerals, they arrange themselves 
into superficial or pre-axial and deep or post-axial divisions, 
which pass, respectively, to the tibial and fibular edges of the 
femur ; but the two are not so separated at their origin by the 
intervention of the prolonged portion which forms the flexor 
of the leg, as the two divisions of the coraco-humerals are 
separated by the prolonged portion of them which forms the 
biceps flexor of the forearm. 

the scapula than to the hnmeras in the instances in which there is no claTide, 
or when it extends beyond that bone. 

The pectoralis minor would seem to be conducted as it were to the thorax in 
Mammals by the costo-coracoid ligament which usually separates it, above, from 
the Bubclayius, and which may be, as suggested by Oegenbaur, a renmant or 
representative of the extension of the coracoid to the sternum in ovipara. 

In PteropUB {Joum. Anat. iii. 801) a deep portion of the peotorid, which 
may be the pect. minora arises from the clavicle. In the Babbit, besides being 
connected with both tubercles of the humerus, it spreads over the supra-spinatust 
and is inserted along the whole length of the spine of the scapula. 

^ In the Dog the subscapularU is partly blended with the supra-spinattu^ and 
in the Mole wiu the teres major. 



MUSCLES IN YEBTSBRATE ANIMALS. S47 

The superficial, or pre-axial, division of the pubischio-femo- 
rals comprises muscles arising from the os pubis (which is proba- 
bly the representative of the precoracoid), and from part of the 
surface of the ischium (which is probably the representative of 
the coracoid). Owing to the large relative size and fixity of 
the bones from which they arise, and the heavier work which 
usually devolves upon the hind limb, they are ordinarily larger 
and more numerous than their correspondent the precoraco- 
humercd with its appendages, the suprarspinatvs and the epi- 
coracO'hv/meralis, in the fore limb. 

The pecUneua is perhaps the most constant segment or 
muscle of the division, and it answers in many respects, serially, 
to the precoraco-htmieral. It arises from the os pubis, and not 
unfrequently extends, over the anterior edge, upon the deep 
surface of the bone, much as the preooraco-humeral in Croco- 
dile extends, over the anterior edge of the precoracoid, upon its 
deep surface. We have found the pectineua in Cryptobranch 
spreading over the dorsal surface of the femur, even to its 
fibular sida Usually, however, its insertion is limited to the 
upper part of the tibial line of the femur (the tibial edge of 
Unea aspera). Where the ala, or anterior edge of the ilium, 
grows into prominence, an extension of this pubischio-femonJ 
division spreads upon and occupies the anterior, or deeper, 
surface of the ala, constituting the iUacua intemus^, in the 
same way that an extension of the superficial coraco-humerals 
spreads upon the anterior surface of the spine of the scapula, 
and constitutes the supra-apinoLtua, Behind the pectineus lies 
the adductor mass, which is variously segmented in different 
animals and which is perforated by a vessel that in some 
animals becomes the main vessel of the leg and foot. It 
sometimes (Hippopotamus, Fig. 48, and others) extends upon 
the tibial side of the leg ; and, we shall find presently, it often 
has intimate relations with the flexors of the leg*. 



^ In Sanrians the iliacut int. ranges, like the peetinew in Ciyptobranoh, but 
to lees extent, over the dorsal snrfaoe of the femor, approaching the fibular side. 
In Mammals it is restricted to the tibial side. 

' The adductor mass not unfrequently extends across the popliteal surface of 
the femur to the fibular line of the linea aspera; and a considerable part of it 
is probably derived from factors which, in the fore limb, range themselves with 
the post-udal division of the coraoo-braohials. 



348 PROFESSOR HUMPHRY. 

The yiew jnst given of the morphological relations of the iUaeus 
intemus derives perhaps some confirmation from the fact that in the 
Hippopotamus, and other artio-dacfyles and some other animalH, there 
is an extension into the pelvis of one of the superficial muscles of the 
thigh — ^the ilio^Hibien of Cnvier. At least near the knee this muscle 
lies in the plane of the sa/rtoriua and the grctcUis and seems, in some 
instances, to belong to the one, and in some, to the other of these two. 
In a Hippopotamus' the muscle I am alluding to was, near the knee, 
joined with the gracUia. Ascending, it diverged from it and divided 
into two portions. Of these, one was inserted into the inner edge of 
the brim of the pelvis, blending there with the lower part of the 
psociB pcurvus. lie other portion expanded over the iUctcua interrms: 
the middle part of this was lost in the fascia covering that muscle; 
and the lateral parts, separating fork-like, approached and blended 
with the p8oa8 jxvrvus near its origin, on the one side, and with the 
outer fibres of the Uiobcus itU, on the other side'. The pectineua was 
not distinctly segmented from the adductors in the Hippopotamus. 

A remarkable derivation from this series of muscles, and 
apparently from the pectinem, is the internal rectus of Saurians 
and Birds. In the Cryptobranch we noticed the pectineue 
extending over the dorsal surface and fibular side of the femur; 
and the interruil rectus would seem to be an extension of the 
lowest part of this, over the knee, beneath the quadriceps, 
into continuity with the flexors of the digits, an extension 
similar to that of a portion of the peroneus — ^the p, fotigrtw— over 
the ankle, through the sole, to the metacarpal of the hallux, or 
to that of the tibialis anticus, in Unau, over the inner side of 
the foot, into the flexor of the digits. 

The deep, or post-axial division of the pubischio-femoral 
mass, which is inserted into the fibular edge of the femur and 
the fibular trochanter, is segmented into the quadraitbs femoris, 
the ffemdli, and the obturator extemus and intemus. The 

^ Fig. 48 represents the masoles of the fore and upper port of the thi{^ 
and of the front of the pelTis, in a young Hippopotamus. II. t., iliacas in- 
iemxiB.^Ps, p., psoas parvns. — Ps. w., psoas magnus. — N, the anterior cniral 
nerye.--I. p. t., ilio-pr6tibien. — Ad. L, addactor longas. —Gr., gracilis.— On Uie 
right side, the gracilis and ilio-pr^tibien have been removed, bringing into view 
more fully the psoas magnus, the iliacus intemus and the adductor longus, also 
shewing (Ad. m.), the adductor magnus passing to the tibia, and {S. t!) the 
semitendinosus. — 8. the sartorins. 

' This extension upon the iliacua is described by Oratiolet in his Mhnoire, 
p. 287, as the Sartoriiu, There is, however, a muscular band passing from the 
ant. spine of the ilium over the front of the patella, which appears to represent 
the aartoritu and the teruor vag. femoris; occasionaUy the gracilU itself extends 
over the peetineua, and so comes into relation with the ilio-pritibien above, aa 
this muscle comes into relation with the graciUi below. 



MUSCLES IN VERTEBRATE ANIMALS. 349 

obturator intemtis appears to result from an extension of 
the mass round the posterior edge of the ischium upon its 
internal surface, resembling the extension which we not unfre- 
quently find of the pectineus upon the internal surface of the 
pubes; and the two divisions of the pubischio-femoral mass 
may thus meet on the internal surface of the pubischium. 
This extension of the obt. int corresponds serially with the 
extension of the coraco-humeral mass upon the posterior and 
inner surface of the coracoid, a prolongation of which (p. 346) 
beneath the scapula forms the subscapvlaria and corresponds 
with the prolongation of the obt. int. beneath the ilium. 



Instead of saying that the obturator intemus is formed by an 
extension of the pubischio-femoral mass round the edge of the iscbium 
upon its internal sarface, it would be more correct to say that the 
fibres of the ventral muscle developed upon the inner surface of the 
pubischium blend posteriorly with, and so form an extension of, 
those on the outer surface. We sometimes find that they take a 
different direction, and blend with the muscles in front of the pubis- 
chium. Thus in Saurians tbey join the members of the pre-axiaJ 
division and, uniting with the pectinetia and Uiacus intemus, create 
an extension of those muscles in that direction. Again in Birds they 
take a third course and, converging to the fore part of the obturator 
foramen, run forwards, through it, to the fibular condyle. 

This remark is applicable to other instances in which I speak 
of an extension of a given muscle over any particular territory. That 
extension is commonly effected by tlie fibres which belong to and are 
developed in the territory mentioned blending with the muscle which 
IB extended over it. Accordingly, it will be commonly found that the 
nerve-supply to the so extended part of the muscle is derived from 
the sources which appertain to the region which it covers. I do not 
therefore attribute much importance to the fact of the internal obtu- 
rator muscle being associated in its nerve-supply with the gemelli 
and quadratus, while the external obturator muscle is in the same 
way rather associated with the adductors and the gracilis. 



The SCAPTJLO-HUMERAL and ILIO-FEMORAL muscles are, 
respectively, situated upon the dorsum of the scapula and of the 
ilium, and are, in the main, inserted into that surface of the 
proximal part of the limb, in each case, which is directed dor- 
sally. This, in the primitive piscine condition of the limb 
(see Dog-fish and Ceratodus), is the dorsal surface. In most 
higher animals, however, the radial tubercle and edge of 
VOL. VI. 24 



350 PROF£SS0B HUMPHRY. 

the humerus, and the fibular tubercle and edge of the femur, 
are turned into the position most favourable for their re- 
ception \ 

In the fore limb the scapulo-humeral mass is, in most of the 
lower animals, represented by a single muscle, diffused upon 
the dorsal surface of the fin in Fishes, but in Urodelans and 
Reptiles converging into a tendon implanted into the radial 
edge or tubercle of the humerus and called dorsalis scapulce. 
This in higher animals is commonly segmented into infra- 
spinatus, teres minor, and teres major. It is probable that 
some of its superficial elements, which in certain cases apply 
themselves to the last-named muscles, in other cases apply them- 
selves to and contribute to the formation of the deltoid. Hence 
we find the latter muscle occasionally (Phoca) encroaching upon 
the territory of these muscles and dwarfing them. The infra- 
spinatus and teres minor are sub-deltoid segments. But the 
teres major may rather be regarded as a sub-laUssimus dorsi 
segment, forasmuch as it applies itself often to the under-surface 
of that muscle. At the same time it is more steadily con- 
nected than the lot. d. with the ulnar ridge of the humerus'. 
Both the teres major and the supra-spinatus sometimes (Pig) 
extend over the respective margins of the scapula upon the 
under-surface, and derive fibres which usually appertain to 
the subscapularis. The teres major does so more particularly; 
and its relation to this territoiy is also shewn by the fact that 
it and the hinder part of the subscapularis muscle are some- 
times supplied by the same subscapular nerve. 

The ilio-femoral mass forms the deep gluteus which is some- 
times one muscle, but is often segmented into the gluteus 
msdius and minimus and the piriformis. The last is, in many 
instances, continuous with the gluteus medius. It commonly 
extends upon the under-surface of the ilium and upon the 

^ This I think is the real explanation of that difference in the insertion of those 
oerresponding muscles in the two limbs which has caused so much difficulty in 
determining their homological relations. The yiew is confirmed by the obser- 
vation that the muscles arising from the dorsum of the scapula are not strictly 
tenacious of their connection with the radial tubercle ; some of them at least in 
Birds and Lizards passing to the ulnar tubercle, or the ulnar edge of the 
humerus. 

' In Seine it passes to the ulnar side of the humerus, quite separate from 
the lot. d. In Manis it is very large, blended with the lat, d, at its origin, and 
aeoompaniea it to the inner condyle of humerus, the olecranon and the forearm. 



MUSCLES IN VERTEBRATE ANIMALS. 351 

sacral vertebrse. It lies beneath the gluteus mix. bearing, at 
the origin, a relation to it similar to that of the ter^ major 
to the laL d. ; and the gluteal vessels and nerves pass between 
its anterior edge and the ilium, just as the posterior branches of 
the subscapular vessels and nerves pass between the anterior 
edge of the teres major and the scapula^. 

Thus the deep or girdle portion of the ventro-appendicular muscle in 
each limb divides itself more or less distinctly into sectors correspond- 
ing generally with, and often to some extent blended with, the sectors 
of the superficial stratum of the same muscle. The coraco-humerals 
in the fore limb, and the adductors in the hind limb, lie beneath, 
and are often respectively blended with, the pectorals and the gracilis. 
The dorscUis scapalcB presents similar relations to the trapezio-deltoid, 
and the teres major to the latissimus dorsi; and the same may be said, 
though less markedly, respecting the relations of the other correspond- 
ing muscles in the hind limb, viz., of the deep glutei with the obli- 
quusy sartoritiSf and tensor vagincB/emoriSf and of the pyriformis with 
the gluteus maximus. 



The Flexors and Extensors of the Forearm and Leo 

are in each instance formed, first, by intrinsic muscles, that 
is, muscles arising from tlie humerus and femur; secondly y by 
extensions of the deep ventro-appendicular stratum, that is, 
by muscles arising from the girdle ; and, thirdly, by extensions 
of the superficial ventro-appendicular stratum; and the muscles 
from these three sources are more or less blended. In the 
simple limbs of Fishes they form only one unbroken layer. 
Some amount of segmentation takes place in most other 
animals; but, owing to the simple nature of the movements 
they effect, the complications and varieties in these muscles 
are not very great or numerous. 

The intrinsic flexors are represented by the brachicUis anticus* 

1 See reasons, in Journal of Anat. ▼. 86, for regarding the teres major and 
the pyriformU to be seriaUy homologous. 

• In Hippopotamus the brcuih. ant, is wanting, the space on the outer side of 
the humerus from which it usually arises being occupied by the «*p. longus which 
is large, and acts as simply a flexor. It might almost be described as hraeh, ant. 
inserted into the lower two-thirds of the radius instead of, as usual, into the 
ulna. The blending of the elements of these two into one is remarkable. The 
muscle winds round the back of the humerus to the inner side, and has a broad 

24—2 



852 PROFESSOR HUMPHRY. 

in the fore limb, and by the short portion of the biceps 
— the femoro-fibular — ^in the hind limb. The latter is sometimes 
inserted low down in the leg (Cryptobranch and Cyclothurus)*. 
In other cases it meets and blends with the biceps flexor cruris, 
and is called the 'short' or 'femoral head of the biceps.' 
It is not unfrequently wanting. Both these are usually in- 
serted into the post-axial bones of the limb— the ulna and 
fibula. The intrinsic dorsal or extensor muscles are repre- 
sented by the parts of the triceps and of the quadriceps which 
arise from the humerus and the femur. They, in the fore 
limb, usually converge to the ulna and, in the hind limb, to the 
tibia. 

The deep ventro-appendicular flexor — the flexor derived from 
the girdle — in the fore limb is the biceps, which presents variable 
degrees of continuity with, and segmentation from, the brachialis 
anticus, and which is further brought into continuity with the 
intrinsic series by means of muscular derivations from the 
humerus in the Bird, and not uncommonly, in other animals, 
including Man in whom they occur as a variety. These are 
generally from the ulnar side of the humerus ; while the Irachi- 
alis anticus extends chiefly from the radial side, between the 
deltoid and the triceps. The coracoid origin of the biceps and 
its relation to the coraco-humeral muscles have been already 
(p. 343) described. Those relations were foimd to be very close in 
Cryptobranch (p. 33), and they shew the biceps to be an inter- 
mediate between the coraco-humerals and the brachialis anticus, 
continuous with either or both, and uniting them into one 
group, which extends from the coracoid, along the ulnar and 
palmar surface of the humerus, to the radius and ulna*. When 
the coracoid reaches to the sternum the origin of the biceps 
is broad, expanding, fan-like, as it ascends; and sometimes 



origin beneath the inner tubercle. It thns dwarfs both of the humeral origins 
of the triceps. It is supplied by the radial nerve. 

^ Joum. Anat. vi. 22 and it. 56. 

* They are aU usually supplied by one nerve — the musculo-cutaneous — which 
is the homologue of tibe coracoid nerve of BepUles, and the serial homologue 
of the obturator nerve. The brachialis anticus is sometimes (Pteropus) found 
to be in direct continuity with the coraco-humeral. In Seine the bicfps de- 
rives two factors from the humerus, which occupy the position of the braehialiM 
anticus. They are so named by Biidinger, see p. 35, and Joum, Anat. ii. 301, 
III. 303, IV. 37. 



MUSCLES IN VERTEBRATE ANIMALS. S5S 

(Owl) it shews a tendency to divide into two— an outer, or 
glenoid, and an inner, or coracoid part. The division is com- 
plete in most Mammals; and the coracoid part forms the more 
superficial element of the muscle. Where the coracoid is very 
short, however (Manis), the division may disappear. The biceps 
is inserted into either or both of the bones of the forearm. 

In the hind limb the flexor derived from the girdle, omitting 
for the present the consideration of the bic^ to which I will 
revert, may, as in Cryptobranch, be blended, in its whole 
length, with the adductor mass ; and it is often united with it 
to some extent. It however has rarely, or never, any connec- 
tion with the femoro-fibular muscle (the short portion of the 
biceps). It is in most animals divided, in its whole length, 
into two. Of these the semitendinosus is usually the more 
free from the adductors S is the more superficial, is inserted 
lower down on the tibia, and is the representative of the cora- 
coid part of the biceps hrachii. The other — the semimembra- 
nosus — is the deeper, represents the glenoid part of the biceps 
brachii, and is more connected with the adductors, presenting 
every degree of cohesion to them. It has also, as have the 
adductors, varied relations with the caudo-feTnoral which some- 
times (some Birds') — ^blends with it, sometimes (Cryptobranch) 
runs to the femur quite separate from it, and sometimes (Sau- 
rians) sends a long tendon, separate from the semimembra/nosvs, 
down to the back of the fibular side of the tibia, or to the 
fibula, or to a sesamoid just above the fibula, from which the 
flexors of the foot and digits arise ». 

^ It retains a conneotion with the gracilis in Pteropns. In Seal the aemi- 
tendino8U8 and semimembranosiu remain one mascle, as their serial homologae 
the biceps brachii sometimes does in Mammals. 

' In the Owl the semimembranosus and adductor are one mascle, which is 
joined by the cattdo-femoral, and is partly continaons with the gastrocnemius. 
In the Swan the caudo-femoral joins the adductor^ which is separate from the 
Bemimembranosns and is inserted into the linea aspera only. In the Gnll the 
eaudo-femoral is inserted into the femnr unconnected with either the semi' 
membr. or the adductor, which are separate; and the latter is partly continaoaB 
with the gastrocnemius, 

' In the Dog the semimembranosus, arising from the tuber ischii, divides 
into two nearly equal portions. Of these one is inserted as usual into the tibia ; 
and the other is inserted into the inner side of the lower part of the femur in 
the locality usually occupied by the adductor magnus. It is there blended with 
a tendon derived from the add. longus, and some fibres are inserted into the 
sesamoid of the- inner head of the gastrocnemius. So that the sesamoid behind 
the internal condyle is a meeting-point between the adductors and the inner 



354 PROFESSOR HUMPHRY. 

The intrinsic extensors are represented by the parts of the 
triceps extensor brachii and the quadriceps extensor cruris, 
which arise from the humerus and femur, and which are very 
constant (except in Cryptobranch) and are subject to but little 
variety. The deep ventro-appendicular, or girdle, element is 
furnished by the scapular origin of the triceps, and by the 
rectus femoris or iliac origin of the quadriceps. Both these are 
connected with the dorsal surface of the girdles near the joints 
with the limbs; but the rotation of the fore limb usually 
directs the origin of the triceps to the hinder edge of the 
scapula, and the rotation of the hind limb directs the rectus 
fevwris to the anterior edge of the ilium*. The superficial 
ventro-appendicular element is furnished in the fore limb by 
the latissimus dorsi, which in Cryptobranch is quite lost in the 
triceps and, in other animals, is partly blended with it; though, 
in Man and some Mammals, it passes to the humerus and has 
no connection with the triceps^. In the hind limb this element 

head of the gastrocmmius ; just as the Besamoid behind the external condyle is a 
meeting-point in Lizards between the tendon of the caudo-femoral and the 
outer head of the gastrocnemius. 

Some of the fibres of the tendon of the adductor magnus in ourselves are 
continued into the internal lateral hgament, and so reach the tibia. In Ai and 
Mania some fibres of the semimembranosus join the add. m. above the knee. In 
the Jerboa the two muscles are inseparably united, and are inserted by a con- 
tinuous tendon into the femur, the tibia, and the Bide of the kuee. In the 
Babbit the adductor is inserted into the tibia, as well as into the femur. In the 
Hippopotamus the semimembr. continues in imion with the adductor as far as 
the knee. 

^ In Bats the rotation of the hind limb is the reverse of what is UFual ; and 
in Pteropus I found the rectus arising from behind the acetabulum. Joum, 
Anat., III. 312. 

The triceps in Manis extends in conjunction with the supinator longus, 
upon the dorsum of the scapula and to the spine, lying over the teres and supra- 
spinatus: and, in Batrachiaus and Beptiles, the triceps derives an origin, be- 
hind, from the coracoid, as, in the same animals, the quadriceps derives an 
origin in front from the pubes. In Seine the long portion of the triceps 
divides, one part passing external to the teres major upon the dorsum of the 
scapula, and the other upon the under surface of the shoulder-joint and the 
coracoid. 

In Phoca (Joum. Anat^ ii. 302) a portion of the triceps (dorsi-epitrochlien), 
blended above with the scapular origin, and arising from the angle of the 
scapula, passes by the olecranon, along the ulnar side of the paddle, to the 5th 
digit. It forms a sort of intermediate between the triceps, the teres major, the 
latissimus dorsi, and the costa alaris of Birds, linking them all together and 
being linked by the two last to the ventral muscle. 

For the paraUel between the relations of the triceps to the radial nerve and 
those of the biceps jlexor cruris to the peroneal nerve, see p. 53. 

s The mode in which the lat. d. blends with the triceps in Cryptobranch 
explains its varied relations with that muscle — its passing in some instances on 
one side and in some on the other side of the scapular origin for instance— 
the fact being that its fibres are in some animalB diffused in the triceps ; in 



>ir 



MUSCLES IN VERTEBRATE ANIMALS. 355 

is furnished by the gluteus maxinms, which, in Birds, is blended 
with the quadriceps, and which, in Cryptobranch and other 
Batrachians, does not exist as an element distinct from the 
rectus^. In Mammals it partly expands into the fascia over- 
lying the quadriceps, and partly is inserted into the fibular line 
of the femur. Sometimes it extends upon the outer side of the 
leg to the ankle and foot. 

I have shewn (p. 22) that in Cryptobranch the long, or 
chief, portion of the biceps flexor cruris is a derivative from the 
gluteo-rectus, that is, from the blended, or unsegmented, super- 
ficial and deep ventro-appendicular extensions upon the dorsum 
of the limb ; and that it inclines upon the lateral surface and 
plantar aspect of the limb, so as to acquire a flexor action. 
It often shews traces of this its origin. It retains its con- 
nection with the ilium in Lizards and Birds, being overlapped 
by the gluteo-rectus in the latter. In Mammals also it is often 
continued upon the ilium through the sacro-sciatic ligament. 
Not unfrequently it has more direct connection with that bone; 
and in some it extends, in close relation with the gluteus, upon 
the caudal vertebrsB*. The inclination of this segment from the 
dorsal to the plantar aspect of the limb, and at the same time 
from an extensor to a flexor function, is probably due to the 
direction of the rotation of the limb. Accordingly, there is an 
absence of any distinct representative of it in Bats ; and the 
nearest approach to the formation of a similar muscle in the 
fore limb is presented by the dorsi-epitrochlien — by such a one 
especially as we find in the Seal (see footnote, p. 354). It 

some they oonverge to and penetrate it at one point, in some at another ; and in 
Bome they oonyerge to, and pass on one side of it. 

^ In Birds there is no distinct rectus femoris ; that is to say, this part of the 
quadriceps is not segmented from the gluteus or, as it might he called, gluteO' 
rectus. This is also the case in Urodelans (p. 22) and is paralleled hy the im- 
perfect segmentation of the latxssiTtms dorsi from the scapular origin of the 
triceps in these animals (p. 36). 

' In Orycteropus and Otter, the biceps arises from the ilium, ilio-sacral liga- 
ment and sacrum. In a wild Cat it was joined near the middle hy a slip from 
the caudal vertebrie just behind the gluteus. In a tame Cat this slip, broad at its 
origin, was continued by a thin tendon which passed between the vastus externus 
and the biceps to the fascia on the outer side of the knee. In the Babbit the 
chief origin of the biceps is from the sacral and caudal vertebrsB ; and it ex- 
pands over the fibula and outer side of the leg, as far as the heel, without being 
inserted into the fibula. In Hippopotamus its disposition resembles that in the 
Babbit ; but it is more blended with the gluteus and has an insertion into the 
fibula. In Seal it reaches to and blends with the extensor tendons on the 
doxrom of the foot. 



356 PROFESSOR HUMPHRY. 

seems in many cases to be segmented &om the ghUeus, much in 
the same manner as the dorsi-epitrochlien is segmented from 
the latiasimus dorsu 

Portions of the triceps, at the lower part of the arm, are 
not unfrequently segmented on one or both sides, constituting 
the anconeus exte^mus and intemus. A similar segmentation 
does not take place in the case of the quadriceps. 

A sesamoid is very often developed in the lower end of the qu^td- 
riceps and occasionally (Pteropus) in the lower end of the triceps. In 
the Wombat and a few other Marsupials the usual tibial sesamoid— 
the patella — is absent and a sesamoid — the fabella — is found upon the 
upper end of the fibula. We may connect this with the relation just 
described between the biceps and the quadriceps. The fabella can 
scarcely be regarded as the homologue of the patella, being connected 
with a different bone and a different part of the dorsal muscle. It 
presents more claim to be the serial homologue of the sesamoid in the 
triceps f situated above the olecranon which is the homologue of 
the large process growing from the upper part of the fibula in Mono- 
tremea The connection of the fabella with an outlying jiortion only 
of the dorsal femoral muscle, whereas the supraH)lecranon sesamoid 
is connected with the middle portion of the dorsal humeral muscle, 
is explained by the fact that the greater part of the femoral muscle 
is directed upon the tibia, whereas the greater part of the humeral 
muscle is directed upon the ulna. 

The insertion of the triceps into the ulna while the quadriceps is 
inserted into the tibia has caused, in the mind of some anatomists, an 
unwillingness to admit the serial homological relation of the two 
muscles. The force of this objection, however, diminishes as we learn 
more of the manner in which the points of insertion as well as of the 
origin of muscles are liable to variation and to wander from one 
bone to another. 

To sum up as thus far traced. The dorsal extension of the 
ventral muscle upon the hind limb gives rise to the gluteus 
rrmgnus, the tensor vaginoe femoris and the sartortus, to the 
rectus femaris and the long part of the biceps, to the pyriformis 
and the glutei. The like extension upon the fore limb gives 
rise to the latissimus dorsi and the trapezio-deltoid, the long 
head of the quadriceps with the dorsi-epitrochlien, to the teres 
major and minor and the infraspinatus. The plantar exten- 
sion of the ventral muscle upon the hind limb gives rise to the 
gracilis and the adductors, in eluding the iliacus intemus on the 
one side and the obturators, gemelli and quadratus femoris on the 
other, together with the semitendinosus and semimembranosus. 



MUSCLES IN VEBTEBRATE ANIMALS. 357 

The same upon the fore limb gives rise to the pectorales and the 
coraco-brachials, including the supra-spinatus on the one side 
and the avhacapularia on the other, together with the biceps. 

Middle and Distal Segments of the Limbs. 

The muscles on the ventral (palmar and plantar) and on 
the dorsal surfaces of these segments of the two limbs are, 
in each instance, as we learn from the lowest orders of Verte- 
brates (see Figs. 23, 24, 34 and 36 and Cryptobranch), in their 
primitive state, in one mass which is connected with, that is to 
say, is to some extent continuous with, the prolongations of the 
ventral muscle upon the limb. In animals above Fishes, in all 
of which a uniform plan of segmentation of the limbs is fol- 
lowed, each mass (palmar or plantar and doi*sal) is usually 
attached to the bone of the upper segment of the limb and to 
the several bones of the middle and distal segments. In ac- 
cordance with the feathering manner in which the limbs, es- 
pecially in the lower animals, are pressed upon the ground and 
withdrawn from it, the fibres on the palmar and plantar surfaces 
are directed from without, inwards as well as downwards, take 
their origin chiefly from the post-axial (ulnar or fibular) con- 
dyle and the post-axial side of the limb, and constitute a 
'pronato-fiexor' mass in each limb; while the fibres on the 
dorsal aspect, having a supinato-extensor action, take a similar 
direction, passing downwards and inwards from the post-axial 
side of the limb, and in some instances, in the hind limb, 
from the dorsal part of the post-axial condyle of the femur. 
In the case of the fore limb, however, the ulna, rising into the 
olecranon, commonly shuts off the supinato-extensor mass of 
muscles from the post-axial condyle of the humerus, and causes 
it — ^the deep as well as the superficial layer of it — to spread 
upon the preaxial condyle and the preaxial edge of the hume- 
rus. A certain method of division of these masses into planes 
and sectors is common to them all, and is also common to them 
in most animals ; though there are of course many varieties in 
detail. A part of either of the masses which associates itself 
with one plane or one sector in a certain limb or animal may 
be otherwise disposed in other cases; but the deviations are 



358 PROFESSOR HUMPHRY. 

on the whole fewer and less than might have been anticipated. 
As a general rule each masa is divisible into a superficial and a 
deep stratum; and the superficial stratum is divisible into 
three sectors — two lateral and a median — though the division 
is by no means equally complete in all instances, that is to say, 
in some animals, as Cryptobranch, the segmentation is much 
less advanced than in others. It is usually less distinct in 
the pronator-flexor mass of the hind limb than in the several 
other masses. There is considerable variation in the extent to 
which the several masses are blended with the muscles of the 
upper segments of the limbs and with the extensions of the 
ventral muscles — the ventro-appendicular muscles — upon the 
limbs. The blending of the muscles of the upper segments is 
more observable on the palmar and plantar than on the dorsal 
aspects, because the dorsal projections of the elbow and knee 
commonly interrupt the continuity between the muscles above 
and below those joints. 

Pronato-flexor Masses. 

The superficial stratum of these in Cryptobranch is very imper- 
fectly segmented from the deep strata in both limbs ; and in the 
hind limb it shows very little trace of division into sectors. In 
that limb it is a simple scarcely segmented mass extending 
from the fibular condyle and the fibular side of the limb upon 
the tarsus and digits. Superficially, it is continuous, through 
the medium of the caudo-pedal, with the superficial prolonga- 
tion of the ventral muscle. This connection is in Lizards main- 
tained only by a delicate tendon, and in higher animals does not 
exist; but the extensions of the gluteus and biceps Are often sub- 
'stituted for it. The deeper (caudo-crural and caudo-femoral) 
prolongations of the ventral muscle are directed chiefly to the 
tibial side of the limb and do not commonly connect themselves 
with the pronato-flexor mass. Such a connection is however, as 
already mentioned, established in some Lizards by the descend- 
ing tendon from the caudo-femoral running into the sesamoid 
above the fibula^ from which the flexors of the foot and toes in 
part arise ; and in some Birds there is a similar connection of 
the same muscle with the inner head of the gastrocnemius 



MUSCLES IN VEBTEBEATE ANIMALS. 359 

through the medium of the semimembranosus. In Manis the 
inner head of the gastrocnemius derives fibres from the adduc- 
tor magnvs; and we know that this muscle is sometimes inti- 
mately connected with the caudo-femoral in the animals in 
which the latter is present. Moreover, the rectus intemus in 
Birds and Lizards is continued over the condyles of the femur 
into the fibular origin of the superficial flexor of the digits, 
affording a good illustration of the manner in which, from the 
primitive unstratified unsegmented homogeneous envelope of 
the limb-skeleton, special forms are evolved. 

Continuing the investigation of this superficial stratum of 
the pronato-flexor mass in the hind limb, in the which, as just 
said, the caudo-pedal is blended, we find it in the lower animals 
where the foot is in a plane with the leg extending, for the 
most part, to the digits. As the heel grows out, however, the 
superficial pronato-flexor stratum becomes concentrated upon 
and more or less inteiTupted by it. 

The plantar fascia^ which represents the pedal part of the 
caudo-pedal, may thus become isolated — cut off by the projecting 
08 calcis — from the crural fascia and from the superficial layers of 
the tendo'Achillis and gastrocnemius, which appear to represent 
the crural part of the caudo-pedal. The pedal part of the flexor 
sublimis digitorum may become, in like manner, cut off from the 
crural part, the former acquiring the name of flexor brevis and 
the latter that of plantaris. The rest of the stratum, uniting 
itself with the caudo-pedal and inserted into the os calcis, forms 
the tendo-AchiUis with the gastrocnemius and soUus, The degree 
however of this calcareal interruption of the superficial pronato- 
flexor stratum varies. The tendo-Achillis is sometimes continued 
into the plantar fascia; and the continuity of i\iQ flexor svblimis 
digitorum from the femur to the digits, as well as its connection 
with the plantar fascia, is very frequent*. In ourselves the 
principle of concentration of force upon the heel is carried 
to the utmost. The pedal and crural parts of the stratum 
are quite isolated from one another by the os calcis ; and addi- 

^ In Phoca (Joum. Anat. n. 814). The elements of the fiexor sublimis are 
derived from the plantaris, the gracilis, the flexor profundus and the tibialis 
posticus. In Unaa also it derives a factor from the tibialis posticus. These are 
interesting remnants of the primitive common hasis from which the several 
masolee were segmented! 



860 PROFESSOR HUMPHRT. 

tional power is brought to bear upon the heel by the origin of 
the soleus from the tibia and by the great development of the 
bellies of the gastrocnemitts. 

Thus considered, the gastrocnemius consists of the blended 
factors of the caudo-pedcU and of the superficial layer of the 
pronato-fiexor mass, some of which — the soleus elements — pass 
from the fibula, with occasionally factors from, or a connection 
with, the caudo-femoral. Most of these descend from the fibu- 
lar side of tbe limb. In Amphibians the fibres that do so 
make up the whole muscle ; and there is nothing to correspond 
with, or represent, the inner or tibial head of Birds and Mam- 
mals. This makes its appearance in Saurians as a thin muscular 
band descending from the lower edge and outer surface of the 
superficial plantar mass of the thigh — the part which represents 
the gracilis and semitendinosus — to the tibial side of the surface 
of the gastrocnemius. In Birds the tibial origin is connected 
rather with the portion which forms the semimembranosus or 
the adductor ; and it acquires also a connection with tibial 
condyle of the femur. In Mammals the relation to the muscles 
descending from the thigh is lost, and the connection with the 
femur only remains, supplemented in Man by an origin from 
the tibia which constitutes the inner or tibial portion of the 
soleus. 

The spreading of the gastrocnemius upon the fore part of the tibia 
in Birds, where it in some (Heron and Gall) is partially blended 
with the sartorius, is another illustration of the connection or con- 
tinnity of the flexor with the extensor, or the plantar with the dorsal, 
muscles, and of the encroachment of one group upon the area of the 
other, of which we have seen so many instances. This portion of the 
muscle, though separated from the remainder by the semitendinosus 
passing between it and the internal condyloid head, is supplied by the 
popliteal nerve. 

The blending of the sartorius with the gastrocnemms above men- 
tioned is an interesting example of that continuity of the extensor 
and flexor muscles of the same limb, which is more remarkably 
exemplified, in a deeper stratum, by the continuity of the internal 
rectus with the flexor digitoru/m. Both subeerve the same purpose, 
viz., that of assisting the action of the femoral flexors of the foot and 
toes during the condition of forced flexion of the knee which is so 
often and long maintained in Birds, and daring which those flexors 
are much relaxed and less capable of acting upon the digits. 

In the fore limb the superficial stratum of the pronato-flexor 



<. 



MUSCLES IN VERTEBRATE ANIMALS. 361 

mass never acquires an origin from the radial condyle, snch as 
would correspond with the tibial head of the gastrocnemius 
which we find in the hind limb of Birds and Mammals. The 
covering fascia^ strengthened by prolongations from the pectoral 
and biceps, perhaps also from the latissimus dorsi, and some- 
times strengthened by muscular fibres derived from the ulnar 
condyle which constitute the palmaris longus, is the representa- 
tive of the caudo-pedal. The most distinct example of a pro- 
longation of the ventral muscle upon the palmar aspect of the 
fore limb— the most distinct serial correspondent, in short, of 
the caudo-pedal — is, however, as stated p. 819, furnished by the 
costo-alaria of Birds*. 

Forasmuch as the carpus retains in all animals the flattened 
form, there is not here, as in the hind limb, a tendency to con- 
centration upon a heel-bone thrown up in the middle, or an 
interruption of the several muscles by such a process. The 
components of this stratum are, therefore, more equally disposed 
upon the surface as they descend, than those of the hind limb ; 
and they present a more distinct division into ulnar, radial and 
intermediate sectors. Of these the ulnar sector {flexor carpi 
ulnaris*), inserted into the cuneiform (with the pisiform) bone 
which is the homologue of the os calcis, is the chief representa- 
tive of the gastrocnemius and soleus. In the intermediate 
sector, the palmaris I expanding into the palmar fascia, where 
it blends with fibres of the flexor carpi ulnaris, represents the 
superficial part of the gastrocnemius; and the flexor suhlimis 
digitorum* represents the plantaris and the flexor brevis 

' I have said that the cotto-alarU is to some extent the serial representative of 
the caudo-pedal, and that the palmaris I, is also a like representative of the 
same muscle. An interesting exemplification of the relation of these parts is 
afforded in Phoca by the palmaris U which, instead of arising as usnal from the 
condyle of the hnmems, passes upon the olecranon and blends with a portion of 
the triceps coming from the angle of the scapula, which again is in close con- 
nection with the latissimus dorsi ; and it spreads like the costo-alaris upon the 
ulnar side of the carpns and hand. The similar exemplification in Manis 
afforded by the direct continuity of the latissimus dorsi with the palmar fascia 
and the ^xor subl. dig. has already been pointed out. 

^ The fl. c. u. is not subject to much variety. It sometimes (Unau and 
Cyclothums) spreads across the wrist superficially, meeting an expansion from 
the supinator longus; or (Phoca) it spreads beneath the palmaris L; or it blends 
with the latter muscle or with the annular ligament. In the Proboscis Monkey it 
meets the sup. I.; and the expansion formed by them lies superficial to, and can 
be dissected off from, the fascia of the forearm. 

> The Jl. d, s. is partially interrupted at the carpus in Pteropus ; and the 
palmaris I. in hi. In Seine the fi. d. s. is tendinous at the wrist and expands 



362 PROFESSOR HUMPHRT. 

digitarum which, as already shewn, are the upper and lower 
parts of the^. 8. dig. pedis. In the hind limb, the last-name<l 
muscle becomes, in higher animals, almost pushed out of the 
field, being reduced to the slender plantaris by the increasing* 
gastrocnemius. But in the fore limb its homologue is in the 
ascendant and absorbs a great part of the fibres of the stratum 
at the expense of the flexor carpi ulnaria and also of the 
palmaris I, which latter dwindles or disappears. The tibial 
sector of the superficial stratum in the hind limb shares the 
fate of the other parts of the stratum; indeed it is altogether 
absorbed by the heel muscle. In the fore limb the correspond- 
ing sector is large, is partly inserted along the radius, forming 
the pronator radii teres, and partly into the carpus or meta- 
carpus, forming the ^earor carpi radialis. 

Thus, granting that the plantaris is repi'esented by the flexor 
digitorum suhlimis^ the elements of the stratum which, in* the 
hind limb, are made to converge into the tendo-AchiUis are, in the 
fore limb, divided into the palmaris longus^ the flexor carpi ulnaris 
and the flexor carpi radialis with part of the pronator teres. More- 
over the palmaris /., instead of being interrupted at the carpus, is 
continued into the palmar fascia; just as the homologous part of the 
stratum in the hind limb is continued into the plantar fascia in those 
animals in which the tarsus retains its primary simple, flat, cai*pu8- 
like form. 

I have before (p. 37, 38) pointed out evidences of the continuity 
of the upper (pronator teres) * part of the radial sector of the super- 
ficial stratum with the deepest (pronator quadrcUus) part of the deeper 
stratum ; and the lower or flexor carpi radicUis part of the sector is, 
in most animals above Urodelans, separate from the pronator teres 
and takes a deep course in the hand*. Now we find in the hind limb 
that the whole of the representatives of this sector — the representa- 
tives, that is, of the pronator teres and the flexor carpi radialis— or 

into a second mnscle below the wrist ; thus closely simulating the plantaris and 
ft. d, t, in the ordinary mammalian ^ind foot. 

^ The ulnar origin of the pronator teres in Man (also in the Chimpanzee and 
Babbit) is an interesting remnant of this continuity. In the Bird (Kite) the 
mnscle is in two distinct parts, both arising from the humerus. Some of the 
branches of the median nerre pass between, some above and some below, the 
two parts. The branches passing between them are distribnted to the fl. c. v. 
and the^. dig. pr. and represent, therefore, part of the nhiar nerve. See dispo- 
sition of the mnscle and its relation to nerves in the Cryptobranoh, p. 87. 

* In the Bird the ftexor carpi radialis ^ like its serial homologue — the tibialis 
posticus — ^lies deep in its whole oonrse. It arises from the ulna only and not 
from the humerus. See varieties of insertion into scaphoid and mets. i., u., 
and III. Joum, Anat.^ rv. 42. In Phoea it is inserted into the scaphoid and Met. i. 
with a slip to Met. ii. ; in the Pig into Met. iix. ; in the Proboscis Monkey into 
Mets. u. and zn. 



/ 

\ 



MUSCLES IN VERTEBBATS AKIMAI^. 363 

Buch of them as are present, viz., the poplUeus and the tibialis poslicuSf 
are situated in a deep level, nearly in the same level with the pronator 
tihice quadratus with which indeed the popliteua is sometimes merged. 
That is to say, the radial sector, though chiefly in a superficial level, is 
continued into a deep level through the medium of certain fibt-es of 
the pronator teres and through the tendon of the flexor carpi radialis; 
whereas the tibial sector, composed of the popliteus and the tibialis 
posticus^ lies througliout in a deep level. 

Indeed, the greatest and most essential features of difference with 
regard to this region in the two limbs ia caused by the fact that the 
flexor carpi radialis, which in the hand lies in as deep a level as does 
the tibialis posticus in the foot and evidently corresponds with it, 
becomes in the forearm paii) of the superficial stratum and is attached 
to the condyle of the humerus; whereas the tibialis posticus continues 
its deep course in the leg and does not reach the femur at all. The 
plane of segmentation of the two miiscles, which is the same in the 
hand and foot, difiers in the forearm and leg. So that, although the 
two tendons might be regarded as serially homologous, the two muscles 
can scarcely be said to be so. Properly viewed, however,- they furnish 
a good illustration of the way in which certain nearly corresponding 
muscles in the two limbs (and the remark applies to muscles of the 
same limb in various animals) are differently segmented from the 
parent mass. They afford further evidence that homological rule is 
not so rigidly followed in development as we may be disposed to think, 
and that we must not be too severe in our attempts to institute homo- 
logical comparison. 

The variation that occurs in the division of the strata and in the 
amount and disposition of the fibres attaching themselves to either stra- 
tum, is also shewn by the fact that in the lower animals the fibres of 
the radial and ulnar sectors are in great part inserted into the radius 
and ulna, so becoming flexors of the forearm upon the arm; and 
superficial prolongations only of them pass on to the carpus ; whereas 
in higher animals the ulnar sector arising partly from the ulna is con- 
tinued in its entirety to the carpus, and the radial sector divides into 
two parts, of which one — the pronator teres — is inserted into the 
radius, and the other — the flexor carpi radialis — is continued to the 
carpus. 

The differences in the two limbs evidently have relation to the 
necessity that exists in higher animals for moving the foot forcibly as a 
whole for the purpose of driving the body onwarda This is accomplish- 
ed by throwing the whole or the greater part of the strength of the 
superficial stratum of the pronato-flexor mass, including the elements 
of the caudo-pedal, upon the projecting lever furnished by the os calcis, 
and by applying a portion of the deeper stratum of the mass (tibialis 
posticus) to the same end as well as to the pronation of the limb ; whereai^ 
in the fore limb the whole of the deep stratum, except the pronator 
quadratus and a considerable part of the superficial stratum, is avail- 
able for the movements of the digits, and so subserves more directly 
to prehension and the requirements of the will. In the hind limb 



364 PROFESSOR HUMPHRY. 

the muscular mass becomes absorbed by the elevator of the heel, and 
in the fore limb it becomes absorbed in the flexors of the digits. 

Not only does the superficial flexor of the digits in the 
fore limb absorb nearly all the elements of the middle sector, 
and push the palmaris I, (the representative of the middle or 
superficial part of the gastrocnemius, including the caudo- 
pedal) nearly or quite out of the field ; but it not unfrequently 
happens that it in turn gives place, in great measure, to the 
deep flexor. Thus it may be reduced (Hatteria), much like its 
homologue in the hind limb of Man, to a mere thin muscle, 
arising near the wrist from a ligament extended between 
the 08 pisiforme and the os naviculare, or (Unau) it may be 
a mere slip from the deep flexor, or it may (Phoca) be im- 
perfectly segmented from the pahnari8\ 

m 

As a prelude to further analysing and reducing the pro- 
nato-flexor masses and entering upon the consideration of 
their deeper strata, we must remember the numerous skeletal 
moveable parts in the distal region of each limb, and must 
conceive an unsegmented chaotic muscular .mass passing from 
above downwards, and dropping fibres upon aaid deriving 

^ In Camivora, where it is nsnally small and often imperfectly Begmented 
from the palmar is y the superficial fibres of its tendons commonly blend with the 
palmar fascia and pass into the pads and tlie skin ; and the deeper fibres form 
the perforated tendons to the phalanges. In the hind limbs of a Dog and of a Cat 
Bome of these fibres to the pad were derived from the Ji. prof, and passed between 
the tendons of the Jl. nibl. Delicate mosciilar slips also passed from the deep 
to the superficial flexor tendons in the sole of these animals. 

In Jerboa the^x. subl. passes to the three middle digits only ; the other digits 
(i and y) receiving dehcate muscular superficial flexors from the supernumerary car- 
pal ossicle which is present in that animal. These, like the same muscular 
Hattena, remind us of the usual disposition of the Jl. subl. in the hind limb. 
In Guinea Pig the superficial flexor sends the greater part of its tendon to join 
the /i prof. The superficial flexor to digit v. is formed on the radial side by a 
tendon from JL prof, and on the ulnar side by a small muscle arising from the 
palmar fascia. In short, the varieties in the segmentation of these muscles are 
very numerous. 

In the Heron and some other birds, the Jl, d, s, is little more than a tendon 
passing, from the int. condyle and the cuneiform bone, to the fascia and the 
proximal phalanx (it passes to the second phalanx in the Swan). In the Jackdaw 
it is a muscular slip from the Jlexor carpi ulnaris. In the Book it is a separate 
smaU muscle arising by a tendon from the int. condyle and passing partly into 
iheJL c. u. and partly to the proximal phalanx of the digit. See varieties in 
disposition and relations between the deep and superficial strata, Joum. Anat. iv. 
43. In Manis the superficial flexor passes to the poUex and is continuous above 
with the latissimui dorsi. 



< 

'S 



MUSCLES IN VERTEBRATE ANIMALS. 365 

fibres from the several skeletal parts after the manner which 
is suggested by the anatomy of the limbs in Fishes and in 
Cryptobranch. 

In the Cryptobranch, segmentation has gone to a certain 
extent. Take the hind limb. First, the superficial fibres pais 
the farthest, extending from the femur, indeed from the tail, 
to the terminal phalanges, and they form a superficial stra- 
tum ; still this stratum is very imperfectly separated from the 
deeper portion of the mass. Secondly (Figs. 9 and 11), a 
band is segmented from the deep part of the mass and is 
directed upon the bases of the digits, forming a flsxor pro- 
fundus digitorum. Thirdly (Fig. 12), several small deep 
bundles are segmented, passing from one skeletal part to 
another near by, so constituting 'intrinsic' muscles — tarso- 
metatarscdes, metatarso-phalangei, phalcmgei and intermetatar- 
sales or interossei. The residuum of the mass constitutes what 
I have called the pronator pedis (Fig. 10). And there is a 
similar arrangement in the fore limb. 

In higher animals the proximal skeletal parts move less 
upon one another ; and \hQ flexor profwndv^ grows at the expense 
of the other components of the deep stratum first mentioned, 
the germs of which are either absorbed or dwarfed by it, 
except those of the interossei. The proruttor pedis also in 
part blends with it, the tarsal portion of its origin forming 
the accessorius\ and the fibular portion of it forming the 
flexor haUvcis. This last joins the deeper surface of the flsxor 
profumdus, and may be destined, as its name implies^ chiefly 



^ In ProbowdB Monk^ the ace€9»oriu$ is united with the tibial flexor {flexor 
prof.) only.